Spermiogenesis and spermatozoon ultrastructure of the dilepidid cestode Molluscotaenia crassiscolex (von Linstow, 1890), an intestinal parasite of the common shrew Sorex araneus

Marigo, A.M., Bâ, C.T. and Miquel, J. 2011. Spermiogenesis and spermatozoon ultrastructure of the dilepidid cestode Molluscotaenia crassiscolex (von Linstow, 1890), an intestinal parasite of the common shrew Sorex araneus. —Acta Zoologica (Stockholm) 92 : 116–125. Spermiogenesis in Molluscotaenia crassiscolex begins with the formation of a differentiation zone containing two centrioles. One of the centrioles develops a flagellum directly into the cytoplasmic extension. The nucleus elongates and later migrates along the spermatid body. During advanced stages of spermiogenesis, a periaxonemal sheath appears in the spermatid. Spermiogenesis finishes with the appearance of a single helicoidal crested body at the base of the spermatid and, finally, the narrowing of the ring of arched membranes causes the detachment of the fully formed spermatozoon. The mature spermatozoon of M. crassiscolex exhibits a partially detached crested body in the anterior region of the spermatozoon, one axoneme, twisted cortical microtubules, a periaxonemal sheath, and a spiralled nucleus. The anterior spermatozoon extremity is characterized by the presence of an electron‐dense apical cone and a single spiralled crested body, which is attached to the sperm cell in the anterior and posterior areas of region I, whereas in the middle area it is partially detached from the cell. This crested body is described for the first time in cestodes. The posterior extremity of the male gamete exhibits only the disorganizing axoneme. Results are discussed and compared particularly with the available ultrastructural data on dilepidids sensu lato.     

Ultrastructure of spermiogenesis and mature spermatozoon of Anonchotaenia globata (von Linstow, 1879) (Cestoda,Cyclophyllidea, Paruterinidae)

Yoneva, A., Georgieva, K., Mizinska, Y., Nikolov, P. N., Georgiev, B. B. and Stoitsova, S. R. 2010. Ultrastructure of spermiogenesis and mature spermatozoon of Anonchotaenia globata (von Linstow, 1879) (Cestoda, Cyclophyllidea, Paruterinidae). — Acta Zoologica (Stockholm) 91 : 184–192 The ultrastructure of spermiogenesis and of the spermatozoon of a species of the family Paruterinidae is described for the first time. The spermiogenesis of Anonchotaenia globata starts with the formation of a differentiation zone with two centrioles associated with thin striated roots. One of the centrioles gives rise to a free flagellum followed by a slight flagellar rotation and a proximodistal fusion of the flagellum with the cytoplasmic protrusion. This pattern corresponds to Type III spermiogenesis in cestodes. The spermatozoon consists of five distinct regions. The anterior extremity possesses an apical cone and a single helically coiled crested body. The cortical microtubules are spirally arranged. The axoneme is surrounded by a periaxonemal sheath and a thin layer of cytoplasm filled with electron‐dense granules in Regions I–V. The periaxonemal sheath is connected with the peripheral microtubules by transverse intracytoplasmic walls in Regions III and IV. The nucleus is spirally coiled around the axoneme. Anonchotaenia globata differs from Dilepididae (where paruterinids have previously been classified) in the type of spermiogenesis, the lack of glycogen inclusions and the presence of intracytoplasmic walls. The pattern of spermiogenesis is similar to that in Metadilepididae and Taeniidae, which are considered phylogenetically close to Paruterinidae.     

Ultrastructural study of spermiogenesis and the spermatozoon in Catenotaenia pusilla, an intestinal parasite of Mus musculus

The ultrastructure of spermiogenesis and the mature spermatozoon in Catenotaenia pusilla (Cestoda: Catenotaeniidae) is described. Spermiogenesis is characterized by the presence of a single axoneme which grows on the outside of a cytoplasmic extension at an angle of 45 degrees. Flagellar rotation and proximodistal fusion are produced in this process. The centrioles lack striated roots and an intercentriolar body. In the mature spermatozoon four different regions are described. The anterior extremity is capped by an apical cone and presents two helical crest-like bodies of unequal length. The axoneme, of the 9 + '1' pattern of the Trepaxonemata, presents a periaxonemal sheath. The cortical microtubules form a spiral pattern at an angle of about 40 degrees to the hypothetical spermatozoon axis. The nucleus is kidney- to horseshoe-shaped in cross section. Granules and proteinaceus walls are not observed in the spermatozoon of C. pusilla.     

Spermiogenesis and spermatozoon ultrastructure of the cestode Mosgovoyia ctenoides (Cyclophyllidea: Anoplocephalidae), an intestinal parasite of Oryctolagus cuniculus (Lagomorpha: Leporidae)

Ultrastructural characters in spermiogenesis and spermatozoa are considered important tools to elucidate the phylogenetic relationships within the Platyhelminthes. In the Anoplocephalidae, ultrastructural data refer to the spermatozoon of 14 species, whereas data on spermiogenesis refer to only 7 species. The present study focused on the spermiogenesis and spermatozoon of the anoplocephalid cestode Mosgovoyia ctenoides, as revealed by transmission electron microscopy. Type IV spermiogenesis was detected, beginning with the formation of a differentiation zone containing 2 centrioles, with a centriolar adjunct and vestigial striated rootlets. Different forms of the latter character have been described in other anoplocephalids. This study supports spermiogenesis of type IV as the most frequent in the Anoplocephalidae and confirms the presence of a centriolar adjunct in yet another type IV spermiogenesis species. The spermatozoon of M. ctenoides possesses 1 axoneme of the 9+ '1' trepaxonematan type, 2 crestlike bodies, dense plates, and granules of electron-dense cytoplasmic material, nucleus, and twisted cortical microtubules. It was again confirmed that the presence of granular material and the absence of both a periaxonemal sheath and intracytoplasmic walls are constant characters in the spermatozoa of all the Anoplocephalinae.     

Ultrastructure of Spermiogenesis and the Spermatozoon of Mathevotaenia herpestis(Cestoda), Intestinal Parasite of Atelerix albiventrisin Senegal

Spermiogenesis in M. herpestisbegins with the formation of a differentiation zone which contains two centrioles associated with an electron–dense, finely granular material. This granular material very quickly becomes striated, a median cytoplasmic extension forms, one of the centrioles becomes laterally oriented in a cytoplasmic bud and the other gives rise to a flagellum. After the migration of the nucleus, a helicoidal crested–like body forms, then the old spermatid separates from the residual cytoplasm. The mature M. herpestisspermatozoon exhibits an apical cone of electron–dense material, a crested–like body and cortical microtubules which are electron–dense centred and spiralized except at their posterior extremity where they are parallel to the spermatozoon axis. The axoneme is of the 9 + ‘1’ pattern. It reaches the posterior extremity of the gamete where the cytoplasm is very electron–dense. The presence of centrioles flanked by ‘striated roots’ has never, to our knowledge, been reported in a platyhelminth. Likewise, a nucleus with an annular cross–section and unevenly distributed electron–dense peri–axonemal material has never been described in a cestod.     

扩张莫尼茨绦虫(圆叶目:裸头科)精细胞分化、精子形成及精子形态

本项研究应用光学显微镜、扫描和透射电子显微镜,观察了扩张莫尼茨绦虫的精细胞分化、精子形成全过程及精子的精细结构。扩张莫尼茨绦虫的精细胞分化过程为：1)初级精原细胞主要发生于幼节的睾丸滤泡中;2)次级精原细胞发生不完全分裂形成16个细胞一簇的初级精母细胞群,以共同的中央细胞质相连;3)初级精母细胞的特征为细胞核中出现联会复合体结构;4)紧接着的第二次成熟分裂,产生64个由中央细胞质相连的细胞核较小的精细胞。精子形成始于精细胞中分化区的形成,成熟精子缺乏线粒体,具有质膜和冠状体、1—4个领域排布的质膜下皮层微管,细胞质中存在电子致密的颗粒状物质,具一个不规则形态的细胞核,具有“9 1”类型的轴丝构造,缺乏轴丝周围鞘。从精子的纵切面上可将精子区分为5个区段(Ⅰ一Ⅴ区)。在精子形成过程中,中心粒基部出现螺旋形小根结构在寄生虫中为首次报导;成熟精子具有游离鞭毛,在绦虫中为首次发现[动物学报49(3)：370—379,2003]。     

Ultrastructure of the spermatozoon of Anomotaenia quelea (Mettrick, 1961) (Cestoda,Cyclophyllidea, Dilepididae), an intestinal parasite of Quelea quelea (Aves,Ploceidae) in Senegal

The mature spermatozoon of Anomotaenia quelea exhibits an apical cone of electron-dense material and two helicoidal crest-like bodies. The apical cone near its base is surrounded by a lucent cytoplasm and a spiraled layer of cortical microtubules. The crest-like bodies are of different lengths, spiraled and make an angle of 30–40° to the hypothetical spermatozoon axis. The axoneme is of the 9 + ‘1’ trepaxonematan pattern and is surrounded by a periaxonemal sheath of electron-dense material. The cytoplasm contains in regions III and IV numerous electron-dense granules situated between the periaxonemal sheath and the cortical microtubules. The posterior extremity of the spermatozoon of A. quelea exhibits a nucleus and a disorganized axoneme and cortical microtubules. This type of posterior extremity of the mature spermatozoon has never been described previously in a Dilepididae. Similarly, two crest-like bodies have not been observed before in a dilepidid cestode.     

Mesocestoides lineatus (Goeze, 1782) (Mesocestoididae): new data on sperm ultrastructure

Spermiogenesis and the ultrastructural characters of the spermatozoon of Mesocestoides lineatus are described by means of transmission electron microscopy, including cytochemical analysis for glycogen. Materials were obtained from a golden hamster (Mesocricetus auratus) after experimental infection with tetrathyridia metacestodes obtained from naturally infected lizards (Anolis carolinensis) from Louisiana. Spermiogenesis in M. lineatus is characterized by the orthogonal growth of a free flagellum, a flagellar rotation, and a proximodistal fusion. The zone of differentiation contains 2 centrioles associated with striated rootlets and a reduced intercentriolar body. The mature spermatozoon of M. lineatus lacks a mitochondrion, and it is characterized by the presence of (1) a single, spiraled, crested body 150 nm thick; (2) a single axoneme of the 9+'1' pattern of trepaxonematan Platyhelminthes; (3) a parallel and reduced row of submembranous cortical microtubules; (4) a spiraled cordon of glycogen granules; and (5) a spiraled nucleus encircling the axoneme.     

Ultrastructural study of spermiogenesis and the spermatozoon of the proteocephalidean cestode Barsonella lafoni de Chambrier et al., 2009, a parasite of the catfish Clarias gariepinus (Burchell, 1822) (Siluriformes, Clariidae)

Spermiogenesis in the proteocephalidean cestode Barsonella lafoni de Chambrier et al., 2009 shows typical characteristics of the type I spermiogenesis. These include the formation of distal cytoplasmic protrusions forming the differentiation zones, lined by cortical microtubules and containing two centrioles. An electron-dense material is present in the apical region of the differentiation zone during the early stages of spermiogenesis. Each centriole is associated to a striated rootlet, being separated by an intercentriolar body. Two free and unequal flagella originate from the centrioles and develop on the lateral sides of the differentiation zone. A median cytoplasmic process is formed between the flagella. Later these flagella rotate, become parallel to the median cytoplasmic process and finally fuse proximodistally with the latter. It is interesting to note that both flagellar growth and rotation are asynchronous. Later, the nucleus enlarges and penetrates into the spermatid body. Finally, the ring of arching membranes is strangled and the young spermatozoon is detached from the residual cytoplasm.The mature spermatozoon presents two axonemes of the 9 + ‘1’ trepaxonematan pattern, crested body, parallel nucleus and cortical microtubules, and glycogen granules. Thus, it corresponds to the type II spermatozoon, described in almost all Proteocephalidea. The anterior extremity of the gamete is characterized by the presence of an apical cone surrounded by the lateral projections of the crested body. An arc formed by some thick and parallel cortical microtubules appears at the level of the centriole. They surround the centriole and later the first axoneme. This arc of electron-dense microtubules disorganizes when the second axoneme appears, and then two parallel rows of thin cortical microtubules are observed. The posterior extremity of the male gamete exhibits some cortical microtubules. This type of posterior extremity has never been described in proteocephalidean cestodes. The ultrastructural features of the spermatozoon/spermiogenesis of the Proteocephalidea species are analyzed and compared.     

Ultrastructure of the spermatozoon of Anomotaenia quelea (Mettrick, 1961) (Cestoda,Cyclophyllidea, Dilepididae), an intestinal parasite of Quelea quelea (Aves,Ploceidae) in Senegal

The mature spermatozoon of Anomotaenia quelea exhibits an apical cone of electron-dense material and two helicoidal crest-like bodies. The apical cone near its base is surrounded by a lucent cytoplasm and a spiraled layer of cortical microtubules. The crest-like bodies are of different lengths, spiraled and make an angle of 30–40° to the hypothetical spermatozoon axis. The axoneme is of the 9 + ‘1’ trepaxonematan pattern and is surrounded by a periaxonemal sheath of electron-dense material. The cytoplasm contains in regions III and IV numerous electron-dense granules situated between the periaxonemal sheath and the cortical microtubules. The posterior extremity of the spermatozoon of A. quelea exhibits a nucleus and a disorganized axoneme and cortical microtubules. This type of posterior extremity of the mature spermatozoon has never been described previously in a Dilepididae. Similarly, two crest-like bodies have not been observed before in a dilepidid cestode.     

Spermiogenesis and ultrastructure of the spermatozoon of the liver fluke Fasciola gigantica Cobbold, 1856 (Digenea: Fasciolidae), a parasite of cattle in Senegal

The present paper describes the spermiogenesis and the ultrastructure of the spermatozoon of Fasciola gigantica, as revealed by transmission electron microscopy. Spermiogenesis in F. gigantica begins with the formation of a differentiation zone containing 2 centrioles with associated striated roots and an intercentriolar body between them. Each centriole develops a flagellum. Proximodistal fusion of these flagella with the median cytoplasmic extension gives rise to the spermatozoon. Spermiogenesis in F. gigantica is characterized by the formation of a dorsolateral cytoplasmic expansion, an external ornamentation of the cell membrane, and spinelike bodies. These 3 structures were also observed in the anterior part of the spermatozoon. Our study describes for the first time the simultaneous presence of dorsolateral cytoplasmic expansion, external ornamentation of the plasma membrane, and spinelike bodies in the spermatozoon of a trematode.     

Spermiogenesis and structure of spermatozoa in the oribatid mite,Hafenrefferia gilvipes (C.L. Koch) (Acari,Oribatida)

The process of spermiogenesis and the structure of spermatozoa in the mite, Hafenrefferia gilvipes (Koch) were studied ultrastructurally. Spermiogenesis was divided into six stages. The spermatids at stage 1 have the usual structure. At stage 2 the structure of the mitochondria and their distribution in the spermatid start to change, leading to the formation of specific mitochondrial derivatives which are subsequently incorporated into the nucleus of the spermatozoon. Parallel to the transformation of mitochondria occurs a reorganization of the nuclear material. The fully formed spermatozoon has a tadpole-like shape, with the cell nucleus located in the distended part of the cell, and containing mitochondrial derivatives in its karyoplasm. Acrosome, flagellum and centrioles are absent. The participation of peripherally distributed microtubules, present in spermatids at stages 4 to 6, in the shaping of the spermatozoon has been suggested.     

Reinvestigation of the ultrastructure of spermiogenesis and the spermatozoon of Hymenolepis nana (Cestoda, Cyclophyllidea), parasite of the small intestine of Rattus rattus.

Spermiogenesis in Hymenolepsis nana begins with the formation of a differentiation zone. This is limited at the front by arched membranes, is surrounded by cortical microtubules associated with 12 crested-like bodies, and contains a single centriole made up of doublets. The distal centriole gives rise to a flagellum that grows at the same pace as the cortical microtubules. Migration of the nucleus takes place after the formation of the flagellum. It is followed by the separation of the old spermatid from the residual cytoplasm. The mature H. nana spermatozoon is filiform and lacks mitochondria. The axoneme, of the 9 + "1" pattern of the Platyhelminthes, does not reach the extremities of the spermatozoon. The nucleus is electron dense and is in close contact with the axoneme around which it coils in a spiral making an angle of 10 degrees to 15 degrees with the spermatozoon axis. The cortical microtubules follow a 10 degrees to 15 degrees helicoidal path along almost their whole length, except at their posterior extremity, where they are parallel to the spermatozoon axis. H. nana is distinguished by the early development of 12 crested-like bodies of different lengths and by the existence of a single centriole in the differentiation zone. Such a high number of crested-like bodies had never previously been reported in a cestode.     

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.     

Ultrastructure of the Spermatozoon of Avitellina centripunctata(Cestoda,Cyclophyllidea), a Parasite of the Small Intestine of Cattle in Senegal

The Avitellina centripunctataspermatozoon is filiform, tapered at both ends and lacks mitochondria. The anterior extremity exhibits an apical cone and a crested–like body 150–200 nm thick. The axoneme is surrounded by a fine layer of lucent cytoplasm and a sheath of electron–dense material. The cytoplasm is slightly electron–dense at the anterior tip of the apical cone and in region IV of the spermatozoon. Over the rest of the gamete it is subdivided into numerous electron–lucent compartments by irregularly spaced walls. The nucleus is electron–dense. It interposes itself between the cortical microtubules which are all electron–dense centred and spiralized along their whole length. An apical zone containing spiralized microtubules, cortical microtubules which are all electron–dense centred and stop before reaching the posterior extremity of the nucleus, as well as the subdivision into cavities by intracytoplasmic proteinaceous material, have never been described before in a cestod. Moreover, the crested–like body of Avitellina centripunctatais the thickest of those which have been described to date in the cestods.     

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.     

Spermiogenesis in the cestode Corallobothrium solidum Fritsch, 1886 (Proteocephalidea: Corallobothriinae)

Spermiogenesis of Corallobothrium solidum Fritsch 1886, has been investigated by transmission electron microscopy. The zone of differentiation contains the two centrioles, each with one thin root, being orientated in the same plane only when a single intercentriolar body (ICB) appears between them. A median cytoplasmic process (MCP) develops distally to the two flagella, which are of unequal length, get longer and rotate towards the MCP. The nucleus penetrates into the spermatid body after the fusion of both flagella with the MCP has started. Flagellar roots occur occasionally in some spermatids. New for the Eucestoda are the following findings: 1. cortical microtubules (CMs) are arranged in two short parallel rows in one‐axoneme region of some spermatids; 2. the crested body of spermatid consists either of electron‐dense tubular elements and interposes itself between CMs, or it is rather homogeneous and situated more peripherally above one continuous semicircle of CMs. The present results support previous data that the type of spermiogenesis in proteocephalideans resembles mostly that observed in tetraphyllideans (Onchobothriidae and Phyllobothriidae), thus supporting the view of a close phylogenetic relationship of tetraphyllidean and proteocephalidean cestodes.     

Spermiogenesis in the Nile tilapia Oreochromis niloticus with notes on a unique pattern of nuclear chromatin condensation

Spermiogenesis in the Nile tilapia, Oreochromis niloticus, was observed ultrastructurally. The process of spermatid differentiation can be divided into six distinct stages based mainly on changes in the nucleus of spermatids. During the latter half of the process, nuclear chromatin condenses progressively to form many dense globules, which ultimately adhere tightly to pack the head of mature spermatozoa. During chromatin condensation the nucleus diminishes in size, and part of the nuclear envelope and nucleoplasm forms a vesicular structure that is finally discarded from the cells together with an associated thin layer of cytoplasm. The spermatozoon comprises a roundish head, a relatively small midpiece, and a relatively short flagellum consisting of the usual 9+2 axoneme. No acrosomal structure is developed during spermiogenesis.     

Ultrastructure of spermiogenesis and the mature spermatozoon of Tetrabothrius erostris loennberg, 1896 (Cestoda, Tetrabothriidae)

The spermiogenesis of Tetrabothrius erostris is characterized by the following events: formation of a differentiation zone containing 2 basal bodies and a pair of rootlets; one of the basal bodies gives rise to a free flagellum, the other induces formation of a flagellar bud; rotation at 90° of the flagellum prior to its fusion with the middle cytoplasmic process of the differentiation zone and partial rotation of the flagellar bud; penetration of the nucleus between the rootlets and appearance of a spur-like protrusion in the differentiation zone; elongation and twisting of the differentiation zone, resulting in twisting of the peripheral microtubules and migration of the nucleus; formation of a crested body; proximal densification of the spermatozoon prior to its detachment from the spermatid rosette. The mature spermatozoon has a single axoneme of 9+“1” type and twisted peripheral microtubules. It consists of 3 portions: a proximal part with a crested body, a middle region rich in β-glycogen, and a distal part containing the nucleus. The pattern of spermiogenesis resembles most closely that in phyllobothriid tetraphyllideans, and probably reflects a relationship of the family Tetrabothriidae with this group.     

Spermiogenesis and fine structure of the spermatozoon in a headstander, Chilodus punctatus (Teleostei, Characiformes, Anostomidae)

The main characteristic features of spermiogenesis in Chilodus punctatus (Characiformes) are rotation of the nucleus, development of a nuclear fossa, which extends as a narrow invagination deep into the nucleus and the way in which flagellum is formed. The chromatin condensation proceeds during the spermiogenesis from heterogeneous through homogenous and granular to a highly compact one present in the mature spermatozoon. Mature Ch. punctatus spermatozoon shows a spherical nucleus, short midpiece and flagellum with lateral fins. The centrioles are in perpendicular arrangement and are located in the deep nuclear fossa, which extends towards the anterior pole of the nucleus. The midpiece contains a few mitochondria, which are separated from the anterior fragment of flagellum by the cytoplasmic channel. Spermiogenesis and spermatozoon ultrastructure conform to the pattern observed in other ostariophysans, but for the first time the presence of lateral fins along flagellum has been documented in a representative of Characiformes.