Accessory Tubules Research Articles

The sperm structure of Clinidium canaliculatum (Costa): A contribution to the systematic position of Rhysodidae (Coleoptera: Carabidae)

The systematic position and the phylogenetic relationship of Rhysodidae members is still debated, with some authors considering the group as a separate family of Adephaga, while for others they could be a subfamily of Carabidae. The group have morphological traits quite different from Carabidae and an aberrant behaviour compared to ground beetles being not predaceous. The sperm ultrastructure of C. canaliculatum was studied comparatively with other species of beetles, Carabidae in particular. The results indicate that the sperm structure of this species is similar to that of the Carabinae species. As in these species, C. canaliculatum has sperm conjugates with an apical conical cap protecting the heads and the initial region of flagella. This sperm appearance is also shared by another species of Rhysodidae, Omoglymmius hamatus. The material of the apical cap consists of an electron-dense material with a peculiar outer net configuration. Many species of Carabidae, however, can present a different type of sperm conjugation, the spermatostyle: a long rod-like structure where the individual sperms have only the most apical part inserted in the cortical area and the flagella are completely free. C. canaliculatum sperm are endowed with a mono-layered acrosome, a nucleus of variable shape along its length, a flagellum consisting of a typical axoneme 9 + 9+2, provided with 16 protofilaments in the tubular wall of accessory tubules, two asymmetric mitochondrial derivatives with the left one larger than the opposite one, and the right accessory body elongated and larger than the opposite one. These sperm characteristics, which are shared also by another member of the group, suggest the demotion of the family Rhysodidae to the subfamily Rhysodinae within Carabidae, a result also supported by recent molecular data.

New Findings on the Sperm Structure of Tenebrionoidea (Insecta, Coleoptera).

Simple SummaryTenebrionoidea, with more than 30,000 described species and 30 currently recognized families, is a superfamily of difficult taxonomy. The aim of this work is to support the basal position of the Mordellidae among the beetle tenebrionoids. They have a low number of sperm cells per cysts, contrary to the more derived families of the group; moreover, their sperm are not distributed in two bundles at the opposite poles of the cysts, as occurs in the higher taxa, but their sperm flagella form a loop in the median region so that sperm nuclei are positioned close to the tail end. The sperm structure of two members of higher families, Oedemeridae and Tenebrionidae, are investigated to confirm the data mentioned above. The sperm looping, which also occurs in the closely related Ripiphoridae, could be the consequence of the growth asynchrony between the cyst size and the sperm length. The Mordellidae sperm are characterized, not by small mitochondrial derivatives and accessory bodies, but by a peculiar stiff and immotile thin flagellar posterior region provided with only accessory tubules.The sperm ultrastructure of a few representative species of Tenebrionoidea was studied. Two species belong to the Mordellidae (Mordellistena brevicauda and Hoshihananomia sp.), one species to Oedemeridae (Oedemera nobilis), and one species to Tenebrionidae (Accanthopus velikensis). It is confirmed that Mordellidae are characterized by the lowest number of spermatozoa per cyst (up to 64), a number shared with Ripiphoridae. In contrast, in the two other families, up to 512 spermatozoa per cyst are observed, the same number present, for example, in Tenebrionidae. Also, as in the other more derived families of tenebrionoids studied so far, during spermatogenesis in O. nobilis and A. velikensis, sperm nuclei are regularly distributed in two sets at opposite poles of the cysts. On the contrary, the Mordellidae species do not exhibit this peculiar process. However, during spermiogenesis, the bundles of sperm bend to form a loop in their median region, quite evident in the Hoshihananomia sp., characterized by long sperm. This process, which also occurs in Ripiphoridae, probably enables individuals to produce long sperm without an increase in testicular volume. The sperm looping could be a consequence of the asynchronous growth between cyst size and sperm length. The sperm ultrastructure of the Mordellidae species reveals that they can be differentiated from other Tenebrionoidea based on the shape and size of some sperm components, such as the accessory bodies and the mitochondrial derivatives. They also show an uncommon stiff and immotile posterior flagellar region provided with only accessory tubules. These results contribute to a better knowledge of the phylogenetic relationship of the basal families of the large group of Tenebrionoidea.

Does dentine mineral change with anatomical location, microscopic site and patient age?

ObjectiveTo determine the effect of patient age (young or mature), anatomical location (shallow/deep and central/peripheral) and microscopic site (intertubular/peritubular) on dentine mineral density, distribution and composition. MethodsExtracted posterior teeth from young (aged 19–20 years, N = 4) and mature (aged 54–77 years, N = 4) subjects were prepared to shallow and deep slices. The dentine surface elemental composition was investigated in a SEM using Backscattered Electron (BSE) micrographs, Energy Dispersive X-ray Spectroscopy, and Integrated Mineral Analysis. Qualitative comparisons and quantitative measures using machine learning were used to analyse the BSE images. Quantitative outcomes were compared using quantile or linear regression models with bootstrapping to account for the multiple measures per sample. Subsequently, a Xenon Plasma Focussed Ion Beam Scanning Electron Microscopy (Xe PFIB-SEM) was used to mill large area (100 µm) cross-sections to investigate morphology through the dentine tubules using high resolution secondary electron micrographs. ResultsWith age, dentine mineral composition remains stable, but density changes with anatomical location and microscopic site. Microscopically, accessory tubules spread into intertubular dentine (ITD) from the main tubule lumens. Within the lumens, mineral deposits form calcospherites in the young that eventually coalesce in mature tubules and branches. The mineral occlusion in mature dentine increases overall ITD density to reflect peritubular dentine (PTD) infiltrate. The ITD observed in micrographs remained consistent for age and observation plane to suggest tubule deposition affects overall dentine density. Mineral density depends on the relative distribution of PTD to ITD that varies with anatomical location. SignificanceAdhesive materials may interact differently within a tooth as well as in different age groups.

The sperm ultrastructure of members of basal Tenebrionoidea (Coleoptera)

The sperm ultrastructure of some beetles of Tenebrionoidea was studied with particular attention to those of the Ripiphoridae, Mordellidae, and Meloidae. These three groups are often thought to form a clade, which is the sister group of the remaining Tenebrionoidea. The testes of the two former families have thinner but longer spermatic cysts containing fewer and longer sperm. Within each cyst all sperm cells have the same orientation, but cross sections showed that the orientation of the axonemes alternate between adjacent cysts, possibly due to the cysts bending on themselves. In both families the sperm has a bilayered acrosome and the flagellum, which shows mitochondrial derivatives starting laterally to the nuclear base, has a typical 9 + 9+2 axoneme with accessory tubules provided with 16 protofilaments in their wall, and well-structured triangular shaped accessory bodies. In Mordellistena sp (Mordellidae) sperm, both mitochondrial derivatives and accessory bodies are somewhat asymmetrical. Moreover, the flagellum shows a very thin and long tail end provided with only accessory tubules. Meloidae species have testes with thicker sperm cysts containing numerous shorter sperm. Within the individual cysts the sperm flagella exhibit an alternating orientation of their axonemes as consequence of a peculiar spermatogenetic process. The flagellar structure is similar to that of the above-mentioned species, but the accessory bodies are not well defined and constituted by fuzzy material. In Mylabris hieracii (Meloidae) sperm, the acrosome is flat with a conspicuous perforatorium and its nucleus has a peculiar quadrangular section. Berberomeloe majalis sperm has a large acrosome with an unusual pentagonal perforatorium. The centriolar structure of Mylabris variabilis shows a complex of dense radial links connecting the microtubular structures to the plasma membrane. These results suggest that Ripiphoridae have a closer relationship with Mordellidae than with Meloidae. These findings are in agreement with results obtained with molecular data.

The sperm ultrastructure of Pytho depressus (Linnaeus, 1767) (Coleoptera, Pythidae)

The sperm ultrastructure of Pytho depressus (Pythidae) is described in this study. The sperm are short cells, about 85−90 μm long, with an acrosome consisting of three layers, a cylindrical nucleus, which at its base has the initial region of two mitochondrial derivatives. The flagellum has two well-developed triangular accessory bodies, and a 9 + 9+2 axonemal pattern with accessory tubules provided with 16 protofilaments in their wall. The structure and shape of the accessory bodies are diagnostic characters within the superfamily. The sperm morphology of P. depressus can be easily distinguished from those of Ripiphoridae, Meloidae and Tenebrionidae. The P. depressus sperm are organized in cysts as in other species of the group but the sperm are not well aligned and show an antiparallel orientation, a feature also observed in other tenebrionids.The phylogenetic implications of the observed sperm features are discussed in the context of comparative sperm ultrastructure of other insect species.

Echinoderes ultraabyssalis sp. nov. from the Kuril-Kamchatka Trench – the first hadal representative of the Kinorhyncha (Kinorhyncha: Cyclorhagida)

A new species of echinoderid kinorhynchs, Echinoderes ultraabyssalis sp. nov. collected from oxidized brown clay at the deepest depression of the Kuril-Kamchatka Trench, North-West Pacific, is described and illustrated using light and electron microscopy. This new representative of the largest kinorhynch genus is characterized by the unique set of spines and tubules and can be easily distinguished from other congeners. The first trunk segment shows remarkably wrinkled posterior margin with parallel cuticular notches; the second trunk segment bears pair of adhesive tubules in both sexes; trunk segment 5 with pair of tubules in lateroventral position; trunk segment 8 with lateral accessory tubules; trunk segment 9 with tubules in laterodorsal position; acicular lateroventral spines on trunk segments 6–9 and two acicular middorsal spines on trunk segments 6 and 8. This species is also well recognized by the presence of tooth-like middorsal protuberance on trunk segment 11. Echinoderes ultraabyssalis sp. nov. constitutes the deepest kinorhynch species described so far (9411–9541 m) and the first hadal representative of the Kinorhyncha in the World Ocean.

Sperm morphology of predatory pirate bugs Amphiareus constrictus and Blaptostethus pallescens (Heteroptera: Anthocoridae) with phylogenetic inferences

The sperm morphology of two predatory bugs Amphiareus constrictus and Blaptostethus pallescens, representing the tribes Dufouriellini and Blasptostethini, respectively, was described using light and transmission electron microscopy. The spermatozoa of Amphiareus constrictus and Blaptostethus pallescens are fine and long, each measuring 216.6μm and 181.0μm in length, of which 37.0μm and 11.6μm, respectively, comprise the nuclei. When stained with DAPI (for DNA), the posterior half of the nucleus in B. pallescens exhibited low fluorescence, while in A. constrictus this feature was observed only in the last 6μm. In both species, as in Heteroptera in general, the spermatozoa have, in the head region, an acrosome and nucleus, and in the flagellar region, an axoneme with 9 accessory tubules, 9 peripheral doublets and 2 central microtubules (9+9+2 microtubules), 2 mitochondrial derivatives (MDs), and a centriolar adjunct in the nucleus-flagellum transition. However, unlike most Heteroptera, in these species, the MDs are asymmetric, and the centriolar adjunct is quite long and encompasses completely the posterior nuclear end and the anterior tips of the MDs. These features are considered as derived, thus supporting the condition derived of Anthocoridae within Cimicomorpha. In addition, several traits of the spermatozoa of these two species easily distinguish one species (and probably a tribe) from the other; for example, the difference of formats in the MDs, and the long anterior projection of the centriolar adjunct parallel to the nucleus in B. pallescens.

Two new species of Dracoderes (Kinorhyncha: Dracoderidae) from the Ryukyu Islands, Japan, with a molecular phylogeny of the genus.

Two new species, Dracoderes snufkini sp. nov. and Dracoderes toyoshioae sp. nov., are described from Okinawa, southern Japan. Diagnostic characters of D. snufkini include: lateroventral tubules on segments 2 and 5; a thick, plump middorsal spine on segments 2 and 9; thick plump paradorsal spines on segments 3-8, alternately laterally displaced; and a ventral primary pectinate fringe on segment 1, with long, wide conspicuous tips. Diagnostic characters of Dracoderes toyoshioae include: a middorsal subcuticular structure on segment 1; paradorsal subcuticular structures on segments 2-9, alternately laterally displaced; paradorsal acicular spines arising from subcuticular structures, at least on segment 5; ventrolateral acicular spines on segment 1; lateral accessory tubules on segment 2; lateral accessory subcuticular structures on segments 2-7; lateroventral tubules on segment 5; and lateroventral subcuticular structures on segments 2-10. Molecular phylogenetic analyses based on 18S rRNA, 28S rRNA and mitochondrial COI sequences indicate that D. abei and D. nidhug are more closely related to one another than either is to D. snufkini or D. toyoshioae.

Spermiogenesis and sperm ultrastructure of Machilontus sp (Insecta: Archaeognatha) with phylogenetic consideration

The sperm structure of the jumping bristletail Machilontus sp has been described. The species shares several sperm characteristics with other genera of the same order Archaeognatha. During late spermiogenesis the spermatid bends at half of its length with the two limbs closely apposed within the same plasma membrane. The sperm has a helicoidal bi-layered acrosome with a filamentous perforatorium and a long nucleus. The elongated flagellum consists of an axoneme with 9 accessory microtubules external to the 9+2, two rows of conventional mitochondria and two accessory bodies. The accessory bodies are located lateral to the axoneme and are probably responsible for the shifting of the accessory tubules in two opposite groups of 5 and 4 tubules, respectively. These sperm characteristics represent common traits of all Archaeognatha.

The sperm structure of Cryptocercus punctulatus Scudder (Blattodea) and sperm evolution in Dictyoptera.

Sperm of the dictyopteran key taxon Cryptocercus punctulatus was examined. It has largely maintained a blattodean groundplan condition, with a three-layered acrosome, an elongate nucleus, a single centriole, a conspicuous centriole adjunct material, two connecting bands (=accessory bodies), and a long functional flagellum with a 9+9+2 axoneme provided with accessory tubules with 16 protofilaments and intertubular material. These sperm characters are shared with several other polyneopterans. The sperm of C. punctulatus is very similar to what is found in Periplaneta americana and species of other groups of roaches, including the sperm of Loboptera decipiens described here for the first time. The general sperm organization here described can be assumed for the groundplan of Insecta and Pterygota. The following evolutionary path can be suggested: after the split between Cryptocercidae and the common ancestor of Isoptera, the typical pattern of sperm formation was altered very distinctly, resulting in a duplication or multiplication (Mastotermitidae) of the centrioles. Mastotermes has maintained a certain sperm motility, but with a very unusual apparatus of multiple flagella with a 9+0 axoneme pattern. After the split into Mastotermitidae and the remaining Isoptera, sperm motility was completely abandoned, and different modifications of sperm components occurred, and even the loss of the sperm flagellum.

The sperm ultrastructure of Stictoleptura cordigera (Füssli, 1775) (Insecta, Coleoptera, Cerambycidae).

The spermatozoa of the longhorn beetles Stictoleptura cordigera were ultrastructurally described in this paper. They have an apical bilayered acrosome, an elongated nucleus, a centriole with star-shape links, two asymmetric mitochondrial derivatives partially crystallized and a 9+9+2 flagellar axoneme with accessory tubules provided with 16 protofilaments in their wall. A centriole adjunct is present and gives rise to two thick laminae as accessory bodies, also asymmetrical, to which two relatively small puff-like structures of different size are connected. These features were previously found in the sperm of the cerambycid Morimus asper. The strict similarity of the cerambycid sperm characters with those of curculionoids indicates a clear phylogenetic relationship between Chrysomeloidea and Curculionoidea.

Comparative morphology of spermatozoa and reproductive systems of zorapteran species from different world regions (Insecta, Zoraptera)

The male and female reproductive apparatus of Zorotypus magnicaudelli (Malaysia), Zorotypus huxleyi (Ecuador) and Zorotypus weidneri (Brazil) were examined and documented in detail. The genital apparatus and sperm of the three species show only minor differences. The testes are larger in Z. magnicaudelli. Z. huxleyi lacks the helical appendage in the accessory glands. A long cuticular flagellum is present in Z. magnicaudelli and in the previously studied Zorotypus caudelli like in several other species, whereas it is absent in Z. weidneri, Z. huxleyi, Zorotypus hubbardi, Zorotypus impolitus and Zorotypus guineensis. Characteristic features of the very similar sperm are the presence of: a) two dense arches above the axoneme; b) a 9 + 9+2 axoneme with detached subtubules A and B of doublets 1 and 6; c) the axonemal end degenerating with enlarging accessory tubules; d) accessory tubules with 17 protofilaments; e) three accessory bodies beneath the axoneme; and f) two mitochondrial derivatives of equal shape. The first characteristic (a) is unknown outside of Zoraptera and possibly autapomorphic. The sperm structure differs distinctly in Z. impolitus and Z. hubbardi, which produce giant sperm and possess a huge spermatheca. The presence of the same sperm type in species either provided with a sclerotized coiled flagellum in males or lacking this structure indicates that a different organization of the genital apparatus does not necessarily affect the sperm structure. The flagellum and its pouch has probably evolved within Zoraptera, but it cannot be excluded that it is a groundplan feature and was reduced several times. The fossil evidence and our findings suggest that distinct modifications in the genital apparatus occurred before the fragmentation of the Gondwanan landmass in the middle Cretaceous.

A new species of Dracoderes (Kinorhyncha: Dracoderidae) from Korea provides further support for a dracoderid-homalorhagid relationship.

A new kinorhynch species, Dracoderes nidhug nov. sp., is described from the East Sea, at 160 m depth, off Uljin, Korea. The new species is the fourth that can be assigned to the genus, and is recognized by the presence of dorsal spines on segments 3 to 9 (alternatingly displaced to more paradorsal positions on segments 3 to 8), subdorsal tubules on segment 2, lateroventral tubules on segments 2, 5 and 10, and lateral accessory tubules on segment 8. The new species shows two longitudinal, intracuticular markings on the ventral side of segment 1, which could be interpret as rudimentary plate joints, corresponding to the articulations found on the sternal plates of segment 1 in species of Pycnophyes and Kinorhynchus. The finding brings further support to a closer relationship between Dracoderes and homalorhagid kinorhynchs.

The male and female reproductive systems of Zorotypus hubbardi Caudell, 1918 (Zoraptera)

Here we present an ultrastructural study of the male and female reproductive systems of Zorotypus hubbardi and compare the findings to those presented in an earlier study. The male reproductive system consists of small testes and thin and short deferent ducts opening into a huge seminal vesicle. At the end of the deferent duct a wiredrawer structure is present which initiates the spermatophore formation. A long ejaculatory duct, originating from the seminal vesicle, receives the secretions of three accessory glands. The copulatory organ is a relatively stout structure consisting of two cuticular claspers connected to a ventral sclerite. The testes contain very large and few germ cells (32 sperm in each cyst) which give rise to large sperm characterized by two giant mitochondrial derivatives, two large accessory bodies, and an axoneme with accessory tubules with 17 protofilaments in their tubular wall. In the seminal vesicle the sperm are joined by a secretion to form an elongate spermatophore. The female system consists of panoistic ovarioles, two lateral oviducts, and a common oviduct which receives the spermathecal duct of a huge spermathecal sac in the terminal part of the vagina. The duct is an anterior prolongation of the sac. Its distal part turns back twisting around its proximal portion. At this level a conspicuous muscle layer gives rise to a valve. The bent spermatophore is hosted in the spermathecal sac, with the sperm heads placed in the proximal part of the spermathecal duct. The opening of the duct is close to the female genital opening. The reproductive systems of Zorotypus caudelli and Z. hubbardi, apart from a distinctly different general organization, also have a different sperm structure: those of the former species are free long-moving cells, while the sperm of Z. hubbardi are giant cells joined in a spermatophore. This allows to hypothesize and discuss a different reproductive behaviour in the two species: monandric in Z. hubbardi and polyandric in Z. caudelli. Apparently different forms of selection have resulted in a very uniform general morphology in Zoraptera, and in highly divergent features related to the reproductive system. The presence of 17 protofilaments in the accessory microtubules of the flagellar axoneme is a potential synapomorphy of Zoraptera and Phasmatodea.

Meristoderes gen. nov., a new kinorhynch genus, with the description of two new species and their implications for echinoderid phylogeny (Kinorhyncha: Cyclorhagida, Echinoderidae)

A new kinorhynch genus, Meristoderes gen. nov., and two new species from Spain and the Solomon Islands, respectively, are described. The new genus is distinguished from all other genera by the first segment consisting of a closed cuticular ring, and the second segment having partial tergosternal junctions, and a superficial midventral fold. This is a new cuticular configuration that may shed light into the phylogenetic relationships of echinoderid kinorhynchs. Meristoderes macracanthus gen. et sp. nov. from the Mediterranean coast of Spain is recognised by the presence of middorsal spines on segments 4, 6 and 8, ventrolateral tubules on segment 2, lateroventral tubules on segment 5, lateroventral spines on segments 6–9, lateral accessory tubules on segment 8, one pair of laterodorsal tubules on segment 10. Meristoderes galatheae sp. nov. from the Solomon Islands is recognized by having a middorsal spine on segment 4 only, ventrolateral tubules on segment 2, lateroventral tubules on segment 5, lateroventral spines on segments 6–9, lateral accessory tubules on segment 8 and subdorsal tubules on segment 10. Both species have a pattern of paraventral perforation site clusters on segments 3–9, with conspicuously long bracteate hairs from the posteriormost perforations sites on the segments 3–7 and 3–6, respectively.The new genus Meristoderes gen. nov. is included into the family Echinoderidae Bütschli, 1876 and appears closely related with the genera Cephalorhyncha Adrianov, 1999 and Echinoderes Claparède, 1863. The new information it provides is discussed to clarify the internal phylogeny of Echinoderidae. The terminology for cuticular characters in the overlapping area between consecutive segments is also standardized.

Sperm structure of some Neuroptera and phylogenetic considerations

Spermatozoa from members of Hemerobiidae, Chrysopidae and Mantispidae (Arthropoda; Hexapoda: Neuroptera) have been examined by electron microscopy. In all species examined, the nucleus is surrounded by a nuclear envelope that in its anterior domain, fans out laterally into one (Chrysopidae) or two wings (Hemerobiidae and Mantispidae). Furthermore, the anterior sperm region is surrounded by external dense material. In Mantispidae, sperm dimorphism with two types of spermatozoa is also confirmed: paraspermatozoa (not fertilizing), provided with giant axoneme and mitochondrial derivatives, and euspermatozoa (fertilizing). Spermatozoa of Chrysopidae and Mantispidae are characterized by the lack of an acrosome while sperm cells of Hemerobiidae are provided with a bilayered acrosome. Spermatozoa from all the investigated species have axonemes of the conventional insect type, with a 9+9+2 microtubular pattern and with accessory tubules provided with 16 protofilaments. In all the examined taxa the intertubular material has the same localization also observed in all other previously analysed Neuroptera. The mitochondrial derivatives and the accessory bodies in the three families are also described. Hemerobiidae are characterized by the presence of a large groove of the plasma membrane along the right side of the anterior sperm region, which results in an eccentric position of the axoneme. Chrysopidae have large mitochondrial derivatives, which encircle the axoneme. The peculiar feature regarding the nuclear envelope was not seen in other members of neuropteroid insects. These data are discussed in the light of the phylogenetic relationships of the taxa examined.

Structure and ultrastructure of spermatozoa of Chrysomya megacephala (Diptera: Calliphoridae)

The spermatozoa of Chrysomya megacephala are similar to those described for other Brachycera. In this species, the spermatozoa are long and thin, measuring about 590 μm in length, of which the head region measures approximately 60 μm. The head includes a monolayered acrosome with electron-lucid material, and the shape of the nucleus, in cross-sections, varies from circular to oval with completely condensed chromatin. The centriole was observed in the zone of flagellar implantation, below the “peg” region. In the region of overlap, the followings structures are observed: nucleus, centriolar adjunct, mitochondrial derivatives and axoneme. The two mitochondrial derivatives are of different lengths but similar diameter. The axoneme is of a conventional insectan type with a 9 + 9 + 2 microtubular arrangement, with accessory tubules flanked by the electron-dense intertubular material. The male internal reproductive tract consists of testis, vas deferens, seminal vesicle, accessory glands and ejaculatory duct.

Ultrastructural studies on euspermatozoa and paraspermatozoa in Mantispidae (Insecta, Neuroptera)

Previous studies have demonstrated the presence of sperm dimorphism in the Mantispidae Perlamantispa perla. We extended the study on several other mantidflies. In all the examined species the occurrence of euspermatozoa (typical) and paraspermatozoa (atypical) was established. The euspermatozoa are characterized by the presence of a cylindrical nucleus surrounded by an envelope that fans out laterally into two thin wings of different length. The acrosome seems to be missing. The nucleus is surrounded by extracellular material. The flagellum is provided with a 9 + 9 + 2 axonemal pattern; the accessory tubules contain 16 protofilaments and the intertubular material has the distribution typical of the taxon. Two elongated accessory bodies flank partially the axoneme and connect this structure with the mitochondrial derivatives. The flagellar axoneme of paraspermatozoa consists of an axoneme and two giant mitochondrial derivatives filled with large globular units. The axoneme exhibits a 9 + 9 + 2 pattern, in which the central 9 + 2 units have a normal structure, in that the microtubular doublets are provided with both dynein arms and radial links. On the contrary, the nine accessory microtubules have a large diameter and their tubular wall consists of 40 protofilaments. This comparative study provided evidences about the uniformity of sperm ultrastructure in Mantispidae. The function of non-fertilizing giant sperm in mantidflies is discussed.

Sperm ultrastructure in Chironomoidea (Insecta, Diptera)

The fine structure of spermatozoa from several species of chironomids, of Culicoides sp. (Ceratopogonidae) and of Odagmia pontina (Simulidae) was studied. A synapomorphic feature, consisting of nine kidney-shaped structures forming the centriole adjunct, was found in the chironomid species. All members of Chironomoidea share a mono-layered acrosome and a flagellar axoneme, provided with accessory tubules with 15 protofilaments in their tubular wall. The axoneme has a 9+9+2 pattern, but in an unidentified species of chironomid, a 9+9+0 model was observed where the central complex and the spokes are missing. Sperm motility is, however, maintained in all the examined species. The spermatozoa of this taxon have the tendency to complete maturation during their progression along the deferent ducts. Thus, in the proximal region of these ducts, they often show remnants of the spermatid cytoplasm.

Sperm structure of Limoniidae and their phylogenetic relationship with Tipulidae ( Diptera, Nematocera)

The sperm ultrastructure of a few species of Limoniidae ( Limonia nigropunctata; L. nubeculosa; Chionea n. sp.; C. alpina; C. lutescens) was studied. The two species of Limonia have a monolayered acrosome with crystallized material, a three-lobed nucleus in cross section, a ring of centriole adjunct material and a flagellum which consists of a 9 + 9 + 1 axoneme and a single mitochondrial derivative. The central axonemal tubule is provided with 15 protofilaments in its tubular wall, while the accessory tubules have 13 protofilaments and are flanked by the electron-dense intertubular material. The three species of Chionea share a monolayered acrosome, a nucleus with two longitudinal grooves, a centriole adjunct material which surrounds the centriole and the initial part of the axoneme. The axoneme is of conventional type, with 9 + 9 + 2 microtubular pattern, with accessory tubules provided with 13 protofilaments and intertubular material. However, in C. lutescens the accessory tubules start with 15 protofilaments and transform into a tubule with 13 protofilaments. These data are discussed in the light of the phylogenetic relationship between Limoniidae and Tipulidae. For this purpose, the sperm ultrastructure of Nephrotoma appendiculata was also considered comparatively.