Internal phylogeny of the Chilopoda (Myriapoda, Arthropoda) using complete 18S rDNA and partial 28S rDNA sequences

The internal phylogeny of the 'myriapod' class Chilopoda is evaluated for 12 species belonging to the five extant centipede orders, using 18S rDNA complete gene sequence and 28S rDNA partial gene sequence data. Equally and differentially weighted parsimony, neighbour-joining and maximum-likelihood were used for phylogenetic reconstruction, and bootstrapping and branch support analyses were performed to evaluate tree topology stability. The results show that the Chilopoda constitute a monophyletic group that is divided into two lines, Notostigmophora (= Scutigeromorpha) and Pleurostigmophora, as found in previous morphological analyses. The Notostigmophora are markedly modified for their epigenic mode of life. The first offshoot of the Pleurostigmophora are the Lithobiomorpha, followed by the Craterostigmomorpha and by the Epimorpha s. str. (= Scolopendromorpha + Geophilomorpha), although strong support for the monophyly of the Epimorpha s. lat. (= Craterostigmomorpha + Epimorpha s. str.) is only found in the differentially weighted parsimony analysis.     

On the centipedes of Yemen

Summary

Six species of Scolopendromorpha and two of Geophilomorpha are recorded from Yemen. All are well known species. The collection contained no Lithobiomorpha or Scutigcromorpha.     

Adding mitochondrial sequence data (16S rRNA and cytochrome c oxidase subunit I) to the phylogeny of centipedes (Myriapoda: Chilopoda): an analysis of morphology and four molecular loci

Relationships within Chilopoda (centipedes) are assessed based on 222 morphological characters, complete 18S rRNA sequences for 70 chilopod terminals, the D3 region of 28S rRNA for 65 terminals, 16S rRNA sequences for 54 terminals and cytochrome c oxidase subunit I sequences for 45 terminals. Morphological and molecular data for seven orders of Diplopoda are used to root cladograms for Chilopoda. Analyses use direct character optimization for 15 gap and substitution models. The Pleurostigmophora and Epimorpha s.l. hypotheses are largely stable to parameter variation for the combined data; the latter clade is formalized as the new taxon Phylactometria. The combined data include parameter sets that support either the monophyly of Epimorpha s.str. (=Scolopendromorpha + Geophilomorpha) or Craterostigmus + Geophilomorpha; the former derives its support from morphology and the nuclear ribosomal genes. Monophyly of Lithobiomorpha and the sister group relationship between Lithobiidae and Henicopidae are stable for morphological and combined data, and are also resolved for the molecular data for 14 of 15 parameter sets. The fundamental split in Scolopendromorpha is between Cryptopidae and Scolopendridae sensu Attems. Blind scolopendromorphs unite as a clade in most molecular and combined analyses, including those that minimize incongruence between data partitions. Geophilomorpha divides into Placodesmata and Adesmata under nine of 15 explored parameter sets.     

Comparative sperm ultrastructure in Nemertea

Although the monophyly of Nemertea is strongly supported by unique morphological characters and results of molecular phylogenetic studies, their ingroup relationships are largely unresolved. To contribute solving this problem we studied sperm ultrastructure of 12 nemertean species that belong to different subtaxa representing the commonly recognized major monophyletic groups. The study yielded a set of 26 characters with an unexpected variation among species of the same genus (Tubulanus and Procephalothrix species), whereas other species varied in metric values or only one character state (Ramphogordius). In some species, the sperm nucleus has grooves (Zygonemertes virescens, Amphiporus imparispinosus) that may be twisted and give a spiral shape to the sperm head (Paranemertes peregrina, Emplectonema gracile). To make the characters from sperm ultrastructure accessible for further phylogenetic analyses, they were coded in a character matrix. Published data for eight species turned out to be sufficiently detailed to be included. Comparative evaluation of available information on the sperm ultrastructure suggests that subtaxa of Heteronemertea and Hoplonemertea are supported as monophyletic by sperm morphology. However, the data do not provide information on the existing contradictions regarding the internal relationships of “Palaeonemertea.” Nevertheless, our study provides evidence that sperm ultrastructure yields numerous potentially informative characters that will be included in upcoming phylogenetic analyses. J. Morphol. 2010. © 2010 Wiley‐Liss, Inc.     

Phylogeny of scolopendromorph centipedes (Chilopoda): morphological analysis featuring characters from the peristomatic area

Alternative schemes for the higher‐level systematics of the centipede order Scolopendromorpha have been established from characters of trunk segmentation, including the segmental position of spiracles, and the presence or absence of eyes. A comparative survey of the preoral chamber by light and scanning electron microscopy contributes 16 new characters of the epipharynx and hypopharynx, sampled from 26 species that represent 20 genera. These new data together with 49 additional morphological characters permit cladistic analysis of the major scolopendromorph groups. The shortest cladograms resolve blind Scolopendromorpha as a basal grade within which a clade now classified as Plutoniuminae + Scolopocryptopidae (supported by unreversed characters from the preoral chamber) is sister to the remaining scolopendromorphs. A unique row of bullet‐shaped sensilla between the labral and clypeal parts of the epipharynx provides a new autapomorphy of the Scolopendromorpha. Either 21 or 23 trunk segments optimize at the base of the Scolopendromorpha but in either case homoplasy is forced on the cladogram. New characters from the epipharynx give additional support for the monophyly of several traditional groupings, including Cryptopinae, Scolopendridae, Otostigmini, and Scolopendrini, and a basal resolution of Asanadini within Scolopendridae. Of the two competing hypotheses for the position of the enigmatic Mediterranean Plutonium zwierleini—being either sister to the cryptopid Theatops or sister to all other Scolopendromorpha—the former hypothesis is strongly supported; spiracles on all trunk segments in Plutonium are homoplastic with the state in Geophilomorpha. Observations on feeding behaviour are needed to illuminate convergence in characters of the epipharynx and mandible in Edentistoma (Otostigminae) and Campylostigmus (Scolopendrini). © The Willi Hennig Society 2008.     

Is Cadlinella ornatissima a chromodorid? Sperm ultrastructure in an enigmatic nudibranch (Opisthobranchia,Mollusca)

Summary

Sperm ultrastructure is described for the nudibranch gastropod Cadlinella ornatissima, type species of the genus Cadlinella (Thiele). Although C. ornatissima exhibits most of the sperm features characteristic of other Opisthobranchia and the Pulmonata (a small, rounded acrosomal vesicle, a complex, helical, mitochondrial derivative—partially paracrystalline, coarse fibres associated with the axoneme), it also possesses a number of previously undescribed and possibly unique features (a longitudinally inrolled acrosomal pedestal, an axial structure within the cavity of the acrosomal pedestal, an electron-dense collar at the anterior region of the acrosomal pedestal, the presence of crystalloid bodies within the glycogen helices of the mitochondrial derivative). To our knowledge this is the first report of crystalloid bodies in mature sperm of any mollusc. Collectively this evidence raises questions concerning the affinities and systematic position of Cadlinella within the Nudibranchia. The peculiar nature of the sperm differences, in comparison with other investigated nudibranchs, suggest that Cadlinella is not easily linked to either the Cadlinidae or Chromodorididae, and should be considered incertae sedis.     

Cauda epididymal spermatozoa of the rufous hare wallaby, Lagorchestes hirsutus (Metatheria, Mammalia) imaged by electron and confocal microscopy

The ultrastructure of mature Lagorchestes hirsutus spermatozoa is described for the first time, revealing unusual aspects of sperm structure in macropodid species. The sperm head is ovoid rather than cuneiform, lacks a ventral nuclear groove and has an acrosomal distribution over approximately 85–90% of its dorsal surface. Immediately adjacent to the nuclear membrane the peripheral nucleoplasm in most spermatozoa form an irregular series of distinctive evaginations previously not described in the spermatozoa of any other marsupial. The midpiece is extremely thickened and short, containing no helical network or peripheral plasma membrane specializations. Axonemal structure is unspecialized with no connecting lamellae; dense outer fibres are closely adherent to axonemal doublets. The sperm morphology of this species is highly aberrant in comparison to other macropod taxa and supports the retention of Lagorchestes as a distinctive genus. In light of this new information, skeletal and serological data should be re‐evaluated to determine the true taxonomic and phylogenetic position of this species.     

Spermatozoon ultrastructure of Xenoturbella bocki (Westblad 1949)

Obst, M., Nakano, H., Bourlat, S.J., Thorndyke, M.C., Telford, M.J., Nyengaard, J.R. and Funch, P. 2011. Spermatozoon ultrastructure of Xenoturbella bocki (Westblad 1949). —Acta Zoologica (Stockholm) 92 : 109–115. Here, we report on the sperm ultrastructure of Xenoturbella bocki (Westblad 1949), which we studied for the first time in detail using light, scanning and transmission electron microscopy. The mature spermatozoa are of the bilaterian primitive type, also called aquasperm and develop as uniflagellate sperm consisting of a round head with distinct mitochondria at the base and a 9+2 flagellum of approximately 42 μm in length. The acrosomal complex consists of a small, round electron translucent acrosomal vesicle and a subacrosomal base. There is no separate midpiece, and the mitochondria surround the proximal and distal centriole in the posterior part of the head. The primitive structure of the spermatozoa suggests that these fertilize the egg by free spawning, probably the ancestral mode of fertilization in early bilaterians. When compared to the spermatozoa of other metazoans, we find that the arrangement of organelles in the Xenoturbella sperm shows similarities to a wide range of protostome and deuterostome taxa and does not seem to indicate any particular phylogenetic relationship.     

An ultrastructural study of the spermatozoa ofEulalia sp. (Phyllodocidae),Lepidonotus sp. (Polynoidae),Lumbrineris sp. (Lumbrineridae) andOwenia fusiformis (Oweniidae)

The ultrastructure of the mature spermatozoa of four polychaetes is described:Eulalia sp. (Phyllodocidae),Lepidonotus sp. (Polynoidae),Lumbrineris sp. (Lumbrineridae) andOwenia fusiformis (Oweniidae). All the sperm show features typical of externally fertilizing sperm in having a rounded nucleus, a short unmodified midpiece, and a simple flagellum with a 9+2 axoneme.Owenia fusiformis andLepidonotus sp. have a nuclear cone extending into the subacrosomal space that may act to present the inner acrosomal membrane to the egg during fertilization. The acrosome ofLumbrineris sp. is flattened and crenulated. The sperm ofEulalia sp. is unusual in having the four mitochondria of the midpiece ensheathed by a membrane. Comparisons are made with other polychaete sperm, and the use of sperm ultrastructure as a taxonomic tool within the Polychaeta is discussed.     

Ultrastructure of spermiogenesis, sperm, and the spermatheca in Terebrasabella heterouncinata (Polychaeta: Sabellidae: Sabellinae)

Abstract. The sabellid polychaete, Terebrasabella heterouncinata, forms burrows in gastropod shells. It is a small, intratubular brooder that breeds semi‐continuously. It has been shown to self‐fertilize, but its reproductive biology suggests that some form of sperm transfer must occur between individuals. To gain an understanding of its fertilization biology, the ultrastructure of spermiogenesis and the sperm in T. heterouncinata was described, and the animal examined for sperm storage structures. Spermiogenesis occurs in clusters of eight spermatids. The mature sperm has an elongate nucleus and a bilaterally symmetrical acrosome with twisted subacrosomal spaces. The midpiece is short, with three crescent‐shaped mitochondria, and forms a tight sheath around the axoneme. A single spermatheca, which opens on the inner ventral part of the crown near the buccal region, is present. It is a simple blind‐ending duct that runs below the ventral nerve cord and is longer than 100 μm. This is the first record of a single spermatheca in Sabellidae. The shape of the sperm and the presence of a spermatheca confirm that individuals of T. heterouncinata produce ent‐aquasperm and would normally cross‐fertilize.     

Male gonads morphology,spermatogenesis and sperm ultrastructure of the seahorse Hippocampus guttulatus (Syngnathidae)

Testes morphology, spermatogenetic process and mature sperm ultrastructure were analysed in Hippocampus guttulatus, using both light and transmission electron microscopy. Both testes were organized in a single large germinal compartment, with a central lumen. Spermatocysts only contained spermatogonia and primary spermatocytes. Inside the testis lumen, together with mature sperm, two types of large mono‐nucleate cells, flagellate and aflagellate, were present. Both types of cells were interpreted as developing germ cells precociously released inside the testis lumen, where their maturation was completed. According to the different morphological features of the nuclei, such as chromatin condensation degree, aspect of the nuclear fossa and others, the flagellate cells were unquestionably developing spermatids. On the contrary, the developmental stage of the aflagellate was more difficult to interpreted. They could be secondary spermatocytes or young spermatids. No dimorphic sperm were recognizable, the only sperm type observed have features typical of the intro‐sperm reports in other syngnathids species. They had a cylindrical head, a short midpiece, characterized by two mitochondrial rings housed inside a cytoplasmic collar, and a long flagellum. These and previous data about the same topic reported on other syngnathids species were compared and discussed.     

Ultrastructural Observations on Fertilization in Dionchus remorae (Platyhelminthes,Monogenea, Dionchidae)

Ultrastructural observations are presented for some of the stages occurring during fertilization in Dionchus remorae (a gill parasite of Echeneis naucrates) and are believed to be the first published concerning a monogenean. Fertilized female germ cells were found in the ovary. Several loops of the spermatozoon were present within the oocyte cytoplasm; the sperm nucleus became electron lucent and the parallel peripheral doublets of the axonemes became increasingly divergent. The cortical granules in the oocyte were not released immediately after penetration by the spermatozoon. The homogeneity apparently found in the oocyte ultrastructure and process of fertilization in the monogeneans and digeneans contrasts with the variety that exists in their sperm ultrastructure.     

Sperm Ultrastructure of Tarsius bancanus(Tarsiidae, Primates): Implications for Primate Phylogeny and the Use of Sperm in Systematics

The ultrastructure of the epididymal sperm of Tarsius bancanus is described. The sperm possess the typical eutherian pattern of a dorsoventrally flattened, ovate sperm head, comprising a nucleus capped by a symmetrical acrosome, and a distinct midpiece and principal piece containing a 9 + 9 + 2 arrangement of outer coarse fibres and microtubules. However, unique features are also present. The overall head length (9 μm) equals the greatest for any primate yet examined, and the subacrosomal space ("perforatorium" or "pseudoperforatorium") tilted at 30° to the sagittal axis of the sperm, is described for the first time for mammals. The acrosome extends only a short distance beyond the length of the nucleus of the mature sperm, and a significant reduction in the acrosome to nuclei ratio appears to occur during the final stages of sperm maturation. In contrast to earlier predictions based on the spermatid of Tarsius syrhicta , the mature spermatozoa of Tarsius shows greatest morphological similarity with the sperm of the Anthropoidea, which have a short symmetrical acrosome, than with the Strepsirhini, which have a relatively long acrosome that can be either symmetrical (Lemuriformes) or asymetrical (Lorisiformes). Four proposed phylogenies of the Primates are assessed using comparative sperm ultrastructure. Placing the Tarsiidae as a sister group to the Lorisidae appears the least likely. The sperm data are consistent with the Tarsiidae being a sister group to the Anthropoidea, to the Strepsirhini, or even to the extant primate groups as a whole. Use of sperm morphology to provide characters in phylogenetic systematics of the primates is discussed, and the principle of "total evidence" is preferred to the common practice of "hanging" sperm on phylogenetic hypotheses based on other evidence. © 1997 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd.     

Ultrastructural observations on the spermatozoa of a species of <Emphasis Type="Italic">Lepidodasys</Emphasis> (Gastrotricha,Macrodasyida)

The ultrastructure of the spermatozoon of a species in the marine gastrotrich genus Lepidodasys is described. The filiform cell is composed of a cork-screw acrosome, a long single mitochondrion surrounded by a helical nucleus, and a flagellum with a 9 × 2 + 2 axonemal arrangement. The structure of the sperm of this species from Denmark appears closely similar to those of the other two species of Lepidodasys studied so far from Italy and Florida (US). Peculiar features (cylindrical nucleus, absence of a periaxonemal sheath) place this genus far from the others in the family Lepidodasyidae. The absence of synapomorphies between Lepidodasys and other genera of Lepidodasyidae suggests that the family is polyphyletic. The sperm ultrastructure fully fits the species of Lepidodasys into the marine order Macrodasyida, with the sperm ground plan of which its sperm shares a number of details.     

Ultrastructure,functional morphology and evolution of recto-canal epidermal glands in Myriapoda

In Chilopoda, solitary epidermal glands are composed of a couple of cells only. These glands are highly abundant on the entire body surface and are distributed throughout the single-layered epidermis. Some authors provided more or less comprehensive observations on the structure of epidermal glands of specific chilopod taxa. However, no information is hitherto available on the ultrastructural diversity of these glands. Furthermore, potential homologies of these chilopod epidermal glands and of their characteristic cellular components remain unknown. Based on our results, we are now able to distinguish two types of epidermal glands in Chilopoda that can be clearly distinguished by their structure and the course of their conducting canal: recto-canal epidermal glands (rceg) and flexo-canal epidermal glands (fceg). In the present paper, we focus on the rceg. We examined the ultrastructural organization of these glands in the head region and on the anterior trunk segments of various representatives of the five extant chilopod orders by light- and electron-microscopy. According to our terminology, rceg consist of up to five different cell types including: a) distal canal cells, b) proximal canal cells, c) intermediary cells, and d) two different types of secretory cells. Intermediary and canal cells form a common conducting canal. The rceg may taxon-specifically differ in relative size and subcellular architecture, but all have the following features in common: 1) a wide distribution on various body regions among all five chilopod subtaxa, 2) the straight, broad and locally dilated conducting canal surrounded by closely packed microvilli or microvilliform infoldings around the apex of the canal cell(s), and 3) the tendency to aggregate to form compound glandular organs of massive size and complexity. Tricellular glandular units established by three different cell types are observed in Scutigeromorpha and Geophilomorpha, whereas four cell types constitute rceg in Lithobiomorpha and Craterostigmomorpha. Five different cell types per glandular unit are found only in Scolopendromorpha. The partial cuticularization of the lower part of the conducting canal formed by the intermediary cell, as found in Chilopoda, differs from the pattern described for equivalent euarthropod epidermal glands, as for instance in Hexapoda. Their wide distribution in Chilopoda and Progoneata makes it likely that tricellular rceg were at least present in the last common ancestor of the Myriapoda. Concerning Chilopoda, the evolution of highly diverse rceg is well explained on the basis of the Pleurostigmophora concept. Glands of the recto-canal type are also found in other arthropods. The paper discusses cases where homology of rceg and also fceg may be assumed beyond Myriapoda and briefly evaluates the potentials and the still-to-be-solved issues prior to use them as an additional character system to reconstruct the phylogeny of the Euarthropoda.     

Two complete mitochondrial genomes in Scolopendra and a comparative analysis of tRNA rearrangements in centipedes

Background

Centipedes are one of the oldest terrestrial arthropods belonging to the sub phylum Myriapoda. With the expansion of our understanding of the application of the two centipedes Scolopendra morsitans and Scolopendra hainanum, belonging to the order Scolopendromorpha, an exhaustive classification was required. Although consensus has been reached on the phylogeny of Chilopoda based on morphological traits, recent analyses based on molecular data exhibited differences in results.

Methods and results

The mitochondrial genome sequences of S. morsitans and S. hainanum were obtained by next-generation sequencing. S. morsitans contains 13 PCGs, two rRNAs, 11 tRNAs, and one CR. whereas S. hainanum contains 12 PCGs, of which ATP8 remains unpredicted, two rRNAs, 14 tRNAs, and one CR. An obvious tRNA rearrangement was found in the genus Scolopendra. S. morsitans exhibited a loss of trnW, trnC, trnI, trnK, trnD, trnA, trnN, trnQ, trnF, trnT, trnS, trnL, and trnV, and a repeat of trnR and trnL. S. hainanum exhibited a loss of trnQ, trnC, trnW, trnI, trnD, trnQ, trnP, and trnV. Phylogenetic analyses of centipedes based on 12 PCGs supported the sister relationship between the orders Geophilomorpha and Lithobiomorpha and a close relationship between Scolopendra dehaani and S. hainanum.

Conclusions

The new mitogenomes determined in this study provide new genomic resources for gene rearrangements and contribute to the understanding of the evolution of gene rearrangement in Chilopoda.

     

Ultrastructure of the male chelicerae of hattena cometis domrow (Acari: Gamasida: Ameroseiidae) functioning as gonopods

The ultrastructure and functional adaptations of male chelicerae in Hattena cometis Domrow are discussed. In particular, as in many other gamasid mites, males of Hattena use the chelicerae, modified as gonopods, to transfer the sperm into the female. For such purpose, a slender process extending from the movable digit, the spermatodactyl, is mainly involved. The spermatodactyl is provided with a sperm transfer duct; in H.cometis, the dorsal surface bent and fused with the ventral surface forms this duct so that the spermatodactyl appears as a cuticular process, connected somehow with the movable digit, and folded on itself to delimit the sperm transfer tube. The organization and ultrastructure of the spermatodactyl are discussed and compared with other gamasid mites studied so far. J. Morphol., 2013. © 2012Wiley Periodicals, Inc.     

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.