Phylogenesis of spermatogenesis in Annulipalpia caddisflies: An ultrastructural analysis on philopotamidae spermiogenesis

Characters currently used in phylogenetic analyses are insufficient for determining the status of several families of the order Trichoptera (caddisflies). Comparative spermatology can contribute to solving this problem. In the suborder Integripalpia, the sperm axoneme displays the pattern of 9 + 2 pairs of parallel microtubules found in many animal species. In the suborder Annulipalpia, however, axonemes bear remarkable aberrations. Consistently, in Chimarra florida (Philopotamidae, Annulipalpia) we found that the number of central microtubules varies from zero to four in axonemes of the spermatids and that spermatozoa lack axonemes. We propose that, similarly, the axoneme pattern became unstable in the Annulipalpia ancestor, from which branched two phylogenetic lines having different kinds of axoneme aberrations: (a) families in which the number of central microtubules of the axoneme exceeds two (e.g., Philopotamidae, Polycentropodidae) and (b) families in which the number of central microtubules of the axoneme does not exceed two (e.g., Hydropsychidae).     

New species of caddisflies (Insecta: Trichoptera) from the Paleogene resins of Europe

Two new species of caddisflies of the families Philopotamidae and Polycentropodidae, Wormaldia pobeda sp. nov. and Nyctiophylax leningrad sp. nov. are described from the Eocene Baltic and Rovno ambers.     

Ultrastructure of sperms in Acoela (Acoelomorpha) and its concordance with molecular systematics

Abstract. The sperms of the Acoela, a group of lower worms, are filiform cells with 2 flagella incorporated into the cell body. Their axonemes can variously have 9+2, 9+1, or 9+0 patterns of microtubules; and singlet microtubules in the cell body can be arranged in axial or cortical positions. An analysis of phylogenetic relationships of acoels based on molecular characters (18S rDNA sequence data) showed that these patterns of microtubules, where known, fell into discrete monophyletic groups. To test this hypothesis, we have expanded the database of sperm characters by examining the ultrastructure of a further 10 species representing 4 acoel families. As expected, the Convolutidae fell into 2 unrelated groups: “small‐bodied convolutids”(Convoluta pulchra, Praeconvoluta tigrina, Pseudaphanostoma smithrii) having 9+2 axonemes and cortical microtubules, and “large‐bodied convolutids” (including Wulguru cuspidata) having 9+0 axonemes and axial microtubules. Also, as expected, a member of the Mecynostomidae (Paedomecynostomum bruneum) has 9+1 axonemes and axial microtubules. Members of a family that appears intermediate by molecular characters, the Otocelididae, significantly have a variety of patterns: axonemes with both 9+2 and 9+0 patterns (Notocelis gullmarensis) or just 9+2 (the other species), and either axial (Philocelis brueggemanni), both axial and cortical (N. gullmarensis) microtubules, or microtubules that bend between axial and cortical positions along the length of the sperm (Otocelis sandara). Members of the Dakuidae (Daku woorimensis) also belong to this intermediate group, having 9+2 axonemes and axial microtubules, while in a fifth otocelidid (Stomatricha hochbergi), sperm characters are like those of the “large‐bodied convolutids” (9+0 axonemes and axial microtubules). Characters of sperm morphology generally support the molecular hypothesis of relationships and confirm a suspected polyphyly of the families Convolutidae, Otocelididae, and Actinoposthiidae.     

Theoretical habitat templets, species traits, and species richness: Trichoptera in the Upper Rhône River and its floodplain

• 1 For Trichoptera occurring in two sites of the Upper Rhône River (France) we examine: (i) relationships among species traits; (ii) habitat utilization of Trichoptera species; (iii) the relationship between species traits and habitat utilization; (iv) trends of species traits in the framework of spatial–temporal habitat variability to test predictions of the habitat templet concept; and (v) trends of species richness in the framework of spatial–temporal habitat variability to test predictions of the patch dynamics concept.
• 2 Of the sixteen species traits selected, twelve have high correlation ratios for the seventy-five species used in this analysis; these traits are related to behavioural, morphological, or physiological aspects. Traits related to reproduction or life cycle have lower correlation ratios.
• 3 An ordination by species traits separates the five main families into three groups: (i) Hydropsychidae and Polycentropodidae; (ii) Hydroptilidae; and (iii) Leptoceridae and Limnephilidae. An ordination of the habitat utilization of the species in ten habitats indicates that the Hydropsychidae occur preferentially in the main channel, Hydroptilidae, Polycentropodidae, and Limnephilidae occur in backwaters or oxbow lakes, and the Leptoceridae are ubiquitous.
• 4 The Hydropsychidae exhibit a relationship between species traits and habitat utilization, i.e. they use similar habitat types with similar species traits. The species traits of the other four families are similar but their habitat utilization is quite different.
• 5 The Hydropsychidae occur in lowest spatial–temporal variability habitats and Limnephilidae in the highest. Therefore, net spinners and filterers are characteristic of habitats with a low spatial–temporal variability, whereas shredders and case makers using plant material are characteristic of habitats with high spatial–temporal variability. The trends in species traits show little agreement with trends predicted from the river habitat templet.
• 6 Trends of species richness in the framework of spatial and temporal variability do not follow the predictions of the patch dynamics concept because richness is similar in all superficial habitats. This implies that each habitat, in spite of large differences in their spatial and temporal variability, offers Trichoptera a similar but limited number of ecological niches.

     

Caddisflies (Trichoptera) of the protected territories of Belarus

The caddisfly fauna of 13 protected areas in Belarus is studied. A total of 97 species from the following 12 families are recorded: Hydroptilidae (6 species), Ecnomidae (1), Polycentropodidae (11), Pshychomyidae (1), Hydropsychidae (5), Phryganeidae (12), Brachycentridae (1), Limnephilidae (41), Sericostomatidae (1), Molannidae (1), Beraeidae (3), and Leptoceridae (14). Polycentropus kingi McLachlan, 1881; Agrypnia picta Kolenati, 1848; Limnephilus sericeus (Say, 1824); L. dispar McLachlan, 1875; L. elegans Curtis, 1834, and Hagenella clathrata (Kolenati, 1848) are rare in Europe. Some of the revealed species are critically endangered in several countries of western Europe. The studied protected areas play an important role in conservation of species diversity and in the distribution of several rare species.     

The dipteran sperm tail: ultrastructural characteristics and phylogenetic considerations

The sperm tail from representatives of several families of Diptera has been examined by high resolution electron microscopy and a computer analysis that improved the visualization of recorded patterns. A considerable variability in sperm tail structure is found within Diptera, and is actually greater than that of any other insect order. The 'generalized insect sperm axoneme'. which is characterized as a 9+9+2 axoneme and by the accessory microtubules having 16 protofilaments, was found only in some dipterans; these are members of Mycetophilidae. From this fact we conclude that Mycetophilidae is likely to be the most primitive extant dipteran group. Another mycetophilid, Boletina , was seen to have accessory tubules with 15 protofilaments as have members of families Dixidae, Chironomidae, Culicidae, and Bibionidae. The last two families have spermatozoa of a type designated as 9+9+'1' there is a central rod rather than two microtubules. We regard this 9+9+'1'pattern with 15 protofilaments to represent a synapomorphic feature. Representatives of the neatoceran families Tipulidae and Trichoceridae have accessory tubules with 13 protofilaments as do examined members of several brachyceran families. Brachycera is hence likely to be derived from the vicinity of the tipulid family. The intertubular material is small in Mycetophilidae and most nematoceran groups, whereas in Tipulidae and Brachycera it is enlarged; here it bridges the space between the accessory tubules and contains various inclusions.     

The pre-impoundment distribution of certain trichoptera larvae in the North Tyne river system (Northern England), with particular reference to current speed

A pre-impoundment survey of the River North Tyne recorded nine species of caseless caddis larvae of the families Hydropsychidae, Rhyacophilidae and Polycentropodidae. 83% of the total numbers collected comprised two species-Hydropsyche siltalai and H. Pellucidula. An analysis was carried out on the distributions of five species with respect to flow rate, from which it appears that the main species-pair H. siltalai, H. pellucidula coexist partly by exhibiting different flow preferences. The possible influence of river impoundment upon flow regimes and invertebrate distributions is briefly discussed.     

New species of caddisflies (Insecta: Trichoptera) from the Rovno amber,Eocene of Ukraine

A new genus and seven new species of the caddisfly families Philopotamidae, Polycentropodidae, Ecnomidae, Hydroptilidae, and Leptoceridae (Wormaldia nasticentia sp. nov., Holocentropus flexiflagrum sp. nov., Electrocyrnus perpusillus gen. et sp. nov., Archaeotinodes reveraverus sp. nov., Agraylea electroscientia sp. nov., Triplectides palaeoslavicus sp. nov., and Leptocerus solifemella sp. nov.) from the Rovno amber (Upper Eocene, Ukraine) are described     

Substructure of the axoneme of pterygote insect spermatozoa: Phylogenetic considerations

Spermatozoa from a great number of insect species were fixed in a tannic acid-containing fixative and the ultrastructure of the flagellar axoneme was examined in a search for apomorphies. Most of the examined species, representing a majority of insect orders. have accessory tubules outside the axoneme (hence a 9 + 9 + 2 pattern), and these consist of 16 protofilaments. Some important apomorphies concern the number of protofilaments in the accessory tubules: 13 (plus 7 inner elements) in Ephemeroptera, 13 in the (elliptic) tubules of Psocoptera + Anoplura + Mallophaga (thus a synapomorphy), 13 in Tipulidae + Brachycera, 15 in the dipteran families Dixidae + Chironomidae (with a 9 + 9 + 2 axoneme) and Culicidae + Bibionidae (with a 9 + 9 + “1” axoneme), 17 in Phasmatodea, and 17–20 in Trichoptera. Other apomorphies concern the appearance of the so-called intertubular material outside the microtubular doublets, the appearance of the interior of the various microtubules, and the loss, in some taxa, of outer or inner dynein arms of both dynein arms. In some cases, the flagellum is completely abnormal; the sperm tail of Thysanoptera, for example, consists of 27 elements of 3 different kinds. The different taxa within orders Diptera and Trichoptera have sperm tail axonemes of different appearances, where those from other orders have a rather uniform appearance. The conclusions that can be drawn from this spermatological study, generally agree with data from classical studies, except with some variations, in some cases.     

Caddisflies from Cenozoic resins of Europe

Analysis of the available data on the findings and taxonomical structure of caddisflies (Insecta, Trichoptera) in the Cenozoic fossil resins of Europe shows that there are four European amber regions (Baltic, Rovno, Saxonian, and Danish) are characterized by a relatively abundant trichopteran fauna, comprising 27 families, 72 genera, and 256 species. These faunas show the dominance of Psychomyioidea (families Polycentropodidae, Psychomyiidae, and Ecnomidae) with Polycentropodidae comprising up to 75% of all records. The amber faunas are second in the dominance of Polycentropodidae only to the terminal Eocene of Florissant (84% of Polycentropodidae). No modern caddisfly species have been found. The amber regions are significantly different in the species composition of Trichoptera although the generic and family structures are similar. Comparison with the modern faunas of Europe shows the absence of advanced Limnephilidae, which are characteristic of the Holocene faunas of Europe, and the rarity of recently abundant Hydropsychidae and Hydroptilidae. The overall composition of amber Trichoptera suggests that it is structurally related to the faunas of Caucasus and Southeastern Asia and might be evidence of seasonally low-contrast (equable) climate in the Late Eocene of Europe.     

New Insects (Insecta: Trichoptera,Reculida, Eoblattida) from the Mesozoic of Asia

New caddisflies, reculid and eoblattid insects from the Mesozoic of Asia are described. Caddisflies of the families Philopotamidae (Mesoviatrix paradoxa gen. et sp. nov. and Kempia piotri gen. et sp. nov.) and Polycentropodidae (Polylongevus eskovi gen. et sp. nov.) from the Upper Jurassic–Lower Cretaceous of the Kempendyai locality in Yakutia are described. A short review and comparison of fossil members in these families are provided. New Gryllones insects, Shurabia tanga sp. nov., Sauk batkenicus gen. et sp. nov. from the Sauk Tan’ga locality (Lower Jurassic of Kyrgyzstan), Say kirgizicus gen. et sp. nov. from the Shurab III locality (Reculida: Geinitziidae; Lower or Middle Jurassic of Kyrgyzstan) Griphopteron iya sp. nov. (Eoblattida: Blattogryllidae) from Iya locality (Middle Jurassic of Russia) are described.     

Sperm ultrastructure and spermiogenesis of Coniopterygidae (Neuroptera, Insecta)

The spermiogenesis and the sperm ultrastructure of several species of Coniopterygidae have been examined. The spermatozoa consist of a three-layered acrosome, an elongated elliptical nucleus, a long flagellum provided with a 9+9+3 axoneme and two mitochondrial derivatives. No accessory bodies were observed. The axoneme exhibits accessory microtubules provided with 13, rather than 16, protofilaments in their tubular wall; the intertubular material is reduced and distributed differently from that observed in other Neuropterida. Sperm axoneme organization supports the isolated position of the family previously proposed on the basis of morphological data.     

ULTRASTRUCTURE OF SWARMERS IN THE LAMINARIALES (PHAEOPHYCEAE). II. SPERM1

The ultrastructure of sperm from 13 species in 11 genera of Laminariales collected in the northeast Pacific Ocean is unique in the brown algae. The sperm are elongate, and possess a nucleus, several mitochondria and two or three chloroplasts, but no eyespot. The anterior flagellum bears mastigonemes on the proximal half of its length; a distal “whiplash” portion lacks mastigonemes and is an extension of only the two central singlet microtubules of the axoneme. A peculiar feature of these sperm is the posterior flagellum, which is longer than the anterior flagellum and tapers distally as the doublet microtubules become singlets and decrease in number. This feature contrasts with the laminarialean zoospore, which possesses a short posterior flagellum with the usual “9 + 2” axoneme. The structure of these sperm differs from that reported for Chorda, the sperm of which resembles a primitive brown algal zoospore. The facts support the concept that Chorda is the most primitive member of the Laminariales.     

Seven new species of caddisflies (Insecta: Trichoptera) from the Rovno amber (Eocene of Ukraine)

Seven new caddisfly species of the families Polycentropodidae, Psychomyiidae, and Lepidostomatidae are described from the Late Eocene Rovno amber: Holocentropus kobodok sp. nov., H. zhiltsovae sp. nov., Plectrocnemia nastigermania sp. nov., P. ucrainica sp. nov., Nyctiophylax terreusbos sp. nov., Lype essentia sp. nov., and Lepidostoma quasitypica sp. nov.     

Ultrastructural analysis of spermatozoa of Korscheltell us lupulinus L. (Lepidoptera : Hepialidae) and Micropterix calthella L. (Lepidoptera : Micropterigidae)

In Korscheltellus lupulinus (Lepidoptera : Hepialidae), both eusperm and parasperm can be found, whereas Micropterix calthella (Lepidoptera : Micropterigidae) produces only eusperm. The eupyrene sperm in K. lupulinus is 250 μm long. Spermatozoa found in K. lupulinus are very similar to those found in higher Lepidoptera, but the eusperm of M. calthella do not possess similar features. As in other Lepidoptera, the axoneme pattern of K. lupulinus is 9 + 9 + 2. Anteriorly, the axoneme is replaced by a fibrous stem similar to an acrosomal structure; posteriorly it becomes progressively uneven. The sperm of K. lupulinus has other original features. The anterior region of the sperm is rigid and contains the nucleus, which has a longitudinal groove adjacent to the axoneme. The 2 mitochondrial derivatives extend from the posterior end of the nucleus. There are no accessory bodies. Many peripheral cytoplasmic vesicles are present. A thin sheath surrounds the cell. It is thick in the parasperm and made of concentric or spiralling lamellae; it does not have any lacinate appendages.The axoneme of the sperm of M. calthella extends over the entire length of the cell and spirals in a helix around the nucleus. The nucleus is also almost as long as the cell. The 2 mitochondrial derivatives, flanked by accessory bodies, follow this spiralling. The axoneme has a 9 + 2 pattern. The plane, which contains the 2 central microtubules, is at an angle with the median plane of the nucleus.     

Ultrastructural and immunocytochemical investigation of acoel sperms with 9 + 1 axoneme structure: new sperm characters for unraveling phylogeny in Acoela

Acoel sperm characters proved useful in deciphering acoel taxonomy. The phylogenetic value of sperm characters in closely related sub-groups or in a monophyletic taxon has not yet been assessed. We have investigated sperm ultrastructure in seven members of the monophyletic taxon Childia sensu (Tekle et al. J Zool Sys Evol Res 43(1):72–90, 2005) and in their closest relatives, the Mecynostomidae (four taxa). All members of Childia examined show little variation in their sperm ultrastructure. The common characters of Childia taxa are: 9 + 1 axoneme structure, the presence of six distal cytoplasmic microtubules in the absence of axial or cortical ones, long nucleus and extensive nucleus–flagella overlap. We have identified a new set of cytoplasmic microtubules lying in the centriolar end of the sperm cell, distal microtubules. The origin and phylogenetic significance of this character is discussed. The types and arrangement of cytoplasmic granules could be used as phylogenetic characters at a low taxonomic level. A loose membrane amorphous core type of granule was found to be a synapomorphy for the following clade within the taxon Childia: C. crassum + C. groenlandica + C. vivipara + C. brachyposthium + C. macroposthium. Sausage shaped granules are plesiomorphic among the taxa examined. The rest of the granule characters were found to be homoplasious. Sperm ultrastructural characters have again proven their concordance with molecular phylogeny. The only morphological synapomorphies known for the sister taxa Childia–Mecynostomidae, in the molecular phylogeny, are characters derived from sperm ultrastructure: distal microtubules arranged in two groups of three microtubules each and a 9 + 1 axoneme structure. The spermatozoa of Childia and Mecynostomidae show 9 + 1 axoneme configuration, seemingly similar to the 9 + ‘1’ axoneme pattern of the Platyhelminthes—Trepaxonemata. Using electron-microscope immunocytochemistry, we have demonstrated that, unlike the central cylinder of trepaxonematans, the central cylinder of the 9 + 1 axonemal pattern in acoels is immunoreactive to tubulin and contains a single central microtubule. Therefore, the 9 + 1 patterns in acoels and trepaxonematans are homoplasious.     

Ten new species of caddisflies (Insecta: Trichoptera) from the Baltic amber

Ten new species of caddisflies of the families Polycentropodidae (nine) and Molannidae (one) are described from the Eocene Baltic amber: Holocentropus peregrinator sp. nov., H. proximorepertus sp. nov., H. dugoi sp. nov., H. fundamentalis sp. nov., H. telergon sp. nov., Plectrocnemia varigoria sp. nov., P. synthesia sp. nov., P. novokshonovi sp. nov., Nyctiophylax valideturbidus sp. nov., and Molanna okraina sp. nov.     

Comparative and functional morphology of wing coupling structures in Trichoptera: Annulipalpia

Several orders of morphologically four‐winged insects have evolved mechanisms that enforce a union between the mesothoracic and metathoracic wings (forewings and hindwings) during the wing beat cycle. Such mechanisms result in a morphologically tetrapterous insect flying as if it were functionally dipterous, and these mechanisms have been described for several insect orders. The caddisfly suborders Annulipalpia and Integripalpia (Trichoptera) each have evolved a wing coupling apparatus, with at least three systems having evolved within the suborder Annulipalpia. The comparative and inferred functional morphology of the putative wing coupling mechanisms is described for the annulipalpian families Hydropsychidae (subfamilies Macronematinae and Hydropsychinae), Polycentropodidae and Ecnomidae, and a novel form‐functional complex putatively involved with at‐rest forewing‐forewing coupling is described for Hydropsychidae: Smicrideinae. It is proposed that the morphology of the wing coupling apparatuses of Hydropsychinae and Macronematinae are apomorphies for those clades. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Anadromous sea lampreys (Petromyzon marinus) are ecosystem engineers in a spawning tributary

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Spermatogenesis of Zaprionus indianus and Zaprionus sepsoides (Diptera,Drosophilidae): Cytochemical,structural and ultrastructural characterization

Zaprionus indianus is a drosophilid native to the Afrotropical region that has colonized South America and exhibits a wide geographical distribution. In contrast, Z. sepsoides is restricted to certain African regions. The two species differ in the size of their testes, which are larger in Z. indianus than in Z. sepsoides. To better understand the biology and the degree of differentiation of these species, the current study evaluated spermatogenesis in males of different ages by conventional staining techniques and ultrastructural analysis. Spermatogenesis and the ultrastructure of spermatozoa were similar in the two species, and the diploid number was confirmed to be 2n = 12. A greater number of spermatozoa were observed in young Z. indianus (1–3 days old) compared to Z. sepsoides males, which showed a higher frequency of cells at the early stages of spermatogenesis. The head of the sperm was strongly marked by silver staining, lacto-acetic orcein and the Feulgen reaction; the P.A.S. reaction revealed glycogen granules in the testes of both species. Both species presented similar arrangement of microtubules (9+9+2), two mitochondrial derivatives of different size and 64 spermatozoa per bundle. Such similarity within the genus Zaprionus with other species of Drosophila, indicates that these structures are conserved in the family Drosophilidae. The differences observed the number and frequency of sperm cells in the early stages of spermatogenesis, between the young males of Z. indianus and Z. sepsoides, are features that may interfere with reproductive success and be related to the invasive potential of Z. indianus.