Muscular architecture of <Emphasis Type="Italic">Milnesium tardigradum </Emphasis>and <Emphasis Type="Italic">Hypsibius </Emphasis>sp. (Eutardigrada,Tardigrada) with some data on <Emphasis Type="Italic">Ramazottius oberhaeuseri</Emphasis>

The architecture of the musculature of the eutardigrade species Milnesium tardigradum Doyère, 1840, Hypsibius sp. and Ramazzottius oberhaeuseri (Doyère in Ann Sci Nat Zool Sér 2(14):269–369, 1840) is investigated by phalloidin staining and confocal laser scanning microscopy. There are methodological problems in staining eutardigrades due to physiological alterations under stress (anhydrobiosis) and due to penetration problems of the cuticle. It is helpful to fix specimens in the state of asphyxy, where animals are stretched following an oxygen shortage in their environment. The musculatures of all three species correspond in their general architecture, but differ in detail, such as in the number of muscles. All muscles are isolated muscle strands. There are on each body side two dorsal and one ventral muscle strands, in addition to a system of dorsoventral, lateral and lateroventral muscles. Seven median ventral attachment points give rise to dorsoventral, ventrolateral and appendage muscles. The appendages receive several muscles originating dorsally and ventrally. The number of muscles and the arrangement differ in each appendage. The fourth appendage shows the greatest differences with a far smaller number of muscles compared to other species. The musculature shows comparably few strict segmental patterns, for example, the musculature of each appendage differs from the other ones. By comparison with literature data on the same species and data of Macrobiotus hufelandi it can be shown that eutardigrades have a roughly comparable muscular architecture, but that there are several differences in detail. Dedicated to Professor Westheide on the occasion of his 70th birthday.     

Functional morphology and neuronal innervation of the prothoracic defence gland in Timema

Timema is the most basal genus of Phasmatodea and the sister group to the remaining stick and leaf-insects (Euphasmatodea). An autapomorphy of all phasmids is the paired prothoracic exocrine defence glands. In this study, the anatomy and innervation of the defence glands in Timema petita and Timema chumash are described and compared with the data on Euphasmatodea. In all phasmids, the glands consist of a cuticular epithelium, a secretory epithelium and muscular fibres that compress the lumen. In Timematodea, the muscular part of the gland is less developed than in Euphasmatodea and the ejection of the defence secretion depends on the dorsal longitudinal neck muscles. On the neuroanatomical level, Timema petita and Timema chumash lack neurons that are involved in the independent contraction of the gland in euphasmids. In both studied species of Timema, neck muscles play an active role in the gland function which is not observed in any other phasmid. Considering the basal position of this genus, this supports the hypothesis that in euphasmids, the muscular part of the gland evolved from the dorsal longitudinal neck muscles. Additionally, the same nerves that innervate the dorsal longitudinal neck muscles in all Polyneoptera also innervate the defence glands in phasmids.     

Muscle formation during embryogenesis of the polychaete Ophryotrocha diadema (Dorvilleidae) – new insights into annelid muscle patterns

Background

The standard textbook information that annelid musculature consists of oligochaete-like outer circular and inner longitudinal muscle-layers has recently been called into question by observations of a variety of complex muscle systems in numerous polychaete taxa. To clarify the ancestral muscle arrangement in this taxon, we compared myogenetic patterns during embryogenesis of Ophryotrocha diadema with available data on oligochaete and polychaete myogenesis. This work addresses the conflicting views on the ground pattern of annelids, and adds to our knowledge of the evolution of lophotrochozoan taxa.

Results

Somatic musculature in Ophryotrocha diadema can be classified into the trunk, prostomial/peristomial, and parapodial muscle complexes. The trunk muscles comprise strong bilateral pairs of distinct dorsal and ventral longitudinal strands. The latter are the first to differentiate during myogenesis. They originate within the peristomium and grow posteriorly through the continuous addition of myocytes. Later, the longitudinal muscles also expand anteriorly and form a complex arrangement of prostomial muscles. Four embryonic parapodia differentiate in an anterior-to-posterior progression, significantly contributing to the somatic musculature. Several diagonal and transverse muscles are present dorsally. Some of the latter are situated external to the longitudinal muscles, which implies they are homologous to the circular muscles of oligochaetes. These circular fibers are only weakly developed, and do not appear to form complete muscle circles.

Conclusion

Comparison of embryonic muscle patterns showed distinct similarities between myogenetic processes in Ophryotrocha diadema and those of oligochaete species, which allows us to relate the diverse adult muscle arrangements of these annelid taxa to each other. These findings provide significant clues for the interpretation of evolutionary changes in annelid musculature.     

Musculature in three species of <Emphasis Type="Italic">Proales</Emphasis> (Monogononta,Rotifera) stained with phalloidin-labeled fluorescent dye

The musculature in the rotifer species Proales daphnicola, P. reinhardti and P. fallaciosa was stained with phalloidin-labeled fluorescent dye and compared using confocal laser scanning microscopy. All three species share several homologous muscle systems, but each systems detailed morphology varies among the species both concerning appearance, number and location. The obtained results were compared with data from other rotifers and it was concluded that the muscles pars coronalis and the corona sphincter probably represent conditions in Ploima or Monogononta, while incomplete circular muscles and dorsal and ventral trunk retractors might be part of the eurotatorian ground pattern.     

On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta)

Friedemann K., Wipfler B., Bradler S. and Beutel R.G. 2011 . On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta). —Acta Zoologica (Stockholm) 00 : 1–16. External and internal head structures of Phyllium siccifolium are described in detail. The findings are compared with conditions found in other phasmatodeans and members of other neopteran lineages. The compiled 125 characters were analysed cladistically. A clade Eukinolabia (Phasmatodea + Embioptera) was confirmed. Synapomorphies of these two taxa are the shift of the origin of M. tentorioparaglossalis to the hind margin of the prementum, the presence of M. tentorioscapalis medialis, and antennal muscles that originate exclusively on the anterior tentorial arms. Within Eukinolabia, the position of Timema remains somewhat ambiguous because of missing anatomical data. However, it was confirmed as sister group of Euphasmatodea in a monophyletic Phasmatodea. Apomorphic groundplan features of Euphasmatodea are salivary ducts with separate external openings, apically rounded glossae, the presence of the galealobulus, and the reduction of the antennifer. The monophyly of Neophasmatidae was confirmed. Autapomorphies are the loss of M. frontobuccalis posterior, the anteriorly or dorsally directed maxillary palps, and the reduction of the mandibular incisivi. The analysis of characters of the head yielded three new autapomorphies of Phylliinae, the presence of a protuberance on the attachment site of the dorsal tentorial arms, dorsoventrally flattened maxillary‐ and labial palps, and possibly the narrow and U‐shaped field of trichomes on the apical part of the galea.     

The phylogenetic placement of Ostropales within Lecanoromycetes (Ascomycota) revisited

Results of molecular studies regarding the phylogenetic placement of the order Ostropales and related taxa within Lecanoromycetes were thus far inconclusive. Some analyses placed the order as sister to the rest of Lecanoromycetes, while others inferred a position nested within Lecanoromycetes. We assembled a data set of 101 species including sequences from nuLSU rDNA, mtSSU rDNA, and the nuclear protein-coding RPB1 for each species to examine the cause of incongruencies in previously published phylogenies. MP, minimum evolution, and Bayesian analyses were performed using the combined three-region data set and the single-gene data sets. The position of Ostropales nested in Lecanoromycetes is confirmed in all single-gene and concatenated analyses, and a placement as sister to the rest of Lecanoromycetes is significantly rejected using two independent methods of alternative topology testing. Acarosporales and related taxa (Acarosporaceae group) are basal in Lecanoromycetes. However, if the these basal taxa are excluded from the analyses, Ostropales appear to be sister to the rest of Lecanoromycetes, suggesting different ingroup rooting as the cause for deviating topologies in previously published phylogenies.     

A new genus and species of monostiliferous hoplonemertean (Nemertea: Enopla: Monostilifera) from Japan

A new genus and species of monostiliferous hoplonemertean, Diopsonemertes acanthocephala gen. et sp. nov., is described from Otsuchi Bay, Japan. Significant anatomical features of the new form include a body wall longitudinal musculature anteriorly divided into inner and outer layers by connective tissue, no pre-cerebral septum, the presence of a thin coat of diagonal muscle fibres between the body wall longitudinal and circular muscle layers in the foregut body region, cephalic retractor muscles derived only from the inner portion of the divided longitudinal muscles and a rhynchocoel more than half the body length.     

Onychophoran‐like myoanatomy of the Cambrian gilled lobopodian Pambdelurion whittingtoni

Arthropods are characterized by a rigid, articulating, exoskeleton operated by a lever‐like system of segmentally arranged, antagonistic muscles. This skeletomuscular system evolved from an unsegmented body wall musculature acting on a hydrostatic skeleton, similar to that of the arthropods’ close relatives, the soft‐bodied onychophorans. Unfortunately, fossil evidence documenting this transition is scarce. Exceptionally‐preserved panarthropods from the Cambrian Lagerstätte of Sirius Passet, Greenland, including the soft‐bodied stem‐arthropod Pambdelurion whittingtoni and the hard‐bodied arthropods Kiisortoqia soperi and Campanamuta mantonae, are unique in preserving extensive musculature. Here we show that Pambdelurion's myoanatomy conforms closely to that of extant onychophorans, with unsegmented dorsal, ventral and longitudinal muscle groups in the trunk, and extrinsic and intrinsic muscles controlling the legs. Pambdelurion also possesses oblique musculature, which has previously been interpreted as an arthropodan characteristic. However, this oblique musculature appears to be confined to the cephalic region and first few body segments, and does not represent a shift towards arthropodan myoanatomy. The Sirius Passet arthropods, Kiisortoqia and Campanamuta, also possess large longitudinal muscles in the trunk, although, unlike Pambdelurion, they are segmentally divided at the tergal boundaries. Thus, the transition towards an arthropodan myoanatomy from a lobopodian ancestor probably involved the division of the peripheral longitudinal muscle into segmented units.     

Histology and Ultrastructure of the Body Wall in the Phoronid Phoronopsis harmeri

The histology and ultrastructure of the body wall in Phoronopsis harmeriwere studied using light microscopy and TEM. The ectoderm epithelium of tentacles, anterior body region, and ampulla consists of monociliary cells. Gram-negative bacteria were found between microvilli, in the protocuticle of the anterior region, and in the ampulla. The epithelium of the posterior body region lacks both monociliary cells and bacteria. The bundles of nerve fibers run between the layer of epithelial cells and basal membrane. The musculature of the body wall comprises circular and longitudinal muscles. The circular muscle fibers are applied to the basal membrane and constitute a solid layer extending almost throughout the length of the body. This pattern is broken in the posterior body region, where there is no solid layer of circular musculature, and the latter is arranged in isolated muscle bands. In the ampullar (terminal) body region, the inversion of circular and longitudinal muscle layers takes place, so that the latter appears to be pressed against the basal membrane. The apical surfaces of longitudinal muscle cells bear cytoplasmic processes; some of the cells have a flagellum. The basal portion of the longitudinal muscle cells forms a cytoplasmic process containing bundles of tonofilaments. The processes of all cells making up the muscle bands are interwoven and anchored to the basal membrane.     

Three‐dimensional reconstruction of the musculature of various life cycle stages of the cycliophoran Symbion americanus

Cycliophora is a very recently described phylum of acoelomate metazoans with a complex life cycle and a phylogenetic position that has been under debate ever since its discovery in 1995. Symbion americanus, which lives attached to the mouthparts of the American lobster, Homarus americanus, represents the second species described for the phylum. Aiming to increase the morphological knowledge about this cryptic clade, the present study describes the muscle arrangement of the feeding stage, the attached Prometheus larva with the dwarf male inside, the free living male, the Pandora larva, and the chordoid larva of S. americanus using actin staining and confocal laser scanning microscopy. 3D reconstructions of the muscular systems are presented. In the feeding stage, circular muscles compose the buccal funnel aperture. In addition, a pair of muscles runs longitudinally in the buccal funnel. A complex sphincter was found just proximally to the anus, and six longitudinal muscles run from the trunk constriction (“neck”) in basal direction. The musculature of the larval stages and the dwarf male is very complex and includes longitudinal muscles that run dorsally and ventrally. In addition, we found dorso‐ventral muscles. The male has a complex posterior muscle apparatus in the vicinity of the penis. In this stage, X‐ and V‐shaped structures were identified on the dorsal and the ventral side, respectively. Pandora and chordoid larvae possess additional circular muscles. We discuss our findings with respect to muscle elements of other metazoan groups and the chordoid larva of Symbion pandora. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Comparative myoanatomy of cycliophoran life cycle stages

The metazoan phylum Cycliophora includes small cryptic epibionts that live attached to the mouthparts of clawed lobsters. The life cycle is complex, with alternating sexual and asexual generations, and involves several sessile and free‐living stages. So far, the morphological and genetic characterization of cycliophorans has been unable to clarify the phylogenetic position of the phylum. In this study, we add new details on the muscular anatomy of the feeding stage, the attached Prometheus larva, the dwarf male, and the female of one of the two hitherto described species, Symbion pandora. The musculature of the feeding stage is composed of myofibers that run longitudinally in the buccal funnel (two fibers) and in the trunk (variable number of fibers). The mouth opening is lined by a myoepithelial ring musculature. A complex myoepithelial sphincter is situated proximal to the anus. In the attached Prometheus larva, three longitudinal sets of myofilaments run dorsally, laterally, and ventrally along the entire anterior‐posterior body axis. The muscular architecture of the dwarf male is complex, especially close to the penis, in the posterior part of the body. An X‐shaped muscle structure is found on the dorsal side, whereas on the ventral side, longitudinal muscles and a V‐shaped muscle structure are present. These muscles are complemented by additional dorsoventral muscles. The mesodermal muscle fibers attach to the cuticle via the epidermis in all life cycle stages studied herein. The musculature of the female is similar to that of the Pandora larva of Symbion americanus and includes dorsoventral muscles and longitudinal muscles that run in the dorsal and ventral body region. Overall, our results reveal striking similarities in the muscular arrangement of the life cycle stages of both Symbion species. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Reconstruction of the musculature of <Emphasis Type="Italic">Magelona</Emphasis> cf. <Emphasis Type="Italic">mirabilis</Emphasis> (Magelonidae) and <Emphasis Type="Italic">Prionospio cirrifera</Emphasis> (Spionidae) (Polychaeta,Annelida) by phalloidin labeling and cLSM

Recent investigations have suggested that a lack of circular muscle fibers may be a common situation rather than a rare exception in polychaetes. As part of a comparative survey of polychaete muscle systems, the F-actin musculature subset of Magelona cf. mirabilis and Prionospio cirrifera were labeled with phalloidin and three-dimensionally analyzed and reconstructed by means of cLSM. Obvious similarities are sublongitudinal lateral, circumbuccal, palp retractor, dominating dorsal longitudinal, perpendicular lateral and ventral transverse muscles. Differences between M. cf. mirabilis and P. cirrifera are: (1) two types of prostomial muscles (transversal and longitudinal) in M. cf. mirabilis versus one type (diagonal) in P. cirrifera; (2) one type of palp muscles (longitudinal) in M. cf. mirabilis versus three types (longitudinal, diagonal, circular) in P. cirrifera; (3) five ventral longitudinal muscles (ventromedian, paramedian, ventral) in M. cf. mirabilis versus four (two paramedian, two ventral) in P. cirrifera. Ventral and lateral transverse fibers are present in the thorax, but absent in the abdomen of M. cf. mirabilis. The triangular lumen of the pharynx in M. cf. mirabilis is surrounded by radial muscle fibers; three sets of pharynx diductors attach to its dorsal side. The unique features of P. cirrifera are one pair of brain muscles and segmentally arranged dorsal transverse muscles, the latter located outside the longitudinal muscles. The transverse lateral muscles are restricted to the sides and lie beneath the longitudinal muscles, a pattern described here for the first time. A true, outer layer of circular fibers is absent in both species of Spionida that were investigated.     

Implications of comparative ventral body wall histology on selection of abdominal surgical approach and closure in 12 species of fish

A ventral midline surgical approach for fish celiotomy is commonly performed in veterinary clinical medicine and research, although the relevant ventral body wall anatomy of many fish species is not well documented. Histological evaluation of tissue samples from the ventral body wall of 12 fish species was performed to provide a reference for surgical approach and closure decisions. The width between muscle bundles running parallel to the long axis and total thickness of tissue layers varied among species. An appreciable space between longitudinal muscles of the ventral body wall and a lack of muscle, vessels and nerves on midline in all species examined supports recommendations of ventral midline incisions to spare important structures. Dense connective tissue consistent with an aponeurosis between musculature along the ventral body wall was not observed in any species evaluated. Connective tissue was concentrated within the dermis of all species evaluated, with an additional layer of collagen along the coelomic membrane in Russian sturgeon Acipenser gueldenstaedtii, koi Cyprinus carpio, goldfish Carassius auratus, black drum Pogonias cromis, black seabass Centropristis striata, tomtate Haemulon aurolineatum and scup Stenotomus caprinus. A sufficiently wide space on ventral midline for practical targeting during the surgical approach is present in A. gueldenstaedtii, C. carpio, striped bass Morone saxatilis, H. aurolineatum, P. cromis, rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta. Sand perch Diplectrum formosum, C. auratus, S. caprinus, grey triggerfish Balistes capriscus and black sea bass Centropristis striata have a negligible space between longitudinal muscles on midline. The variation in ventral body wall structure observed in this study helps inform surgical decision making for celiotomy incision and closure in these species.     

Structure of the mouth tube of a parasitic copepod,Isobranchia appendiculata Heegaard, 1947 (Siphonostomatoida:Lernaeopodidae)

The structure and musculature of the mouth tube of the lernaeopodidIsobranchia appendiculata Heegaard is described in detail. Situated on the median eminence, the mouth tube appears to be lodged inside a median longitudinal groove on the ventral wall of the cephalothorax. The components of the mouth tube, the labrum and the labium are loosely held together along their lateral edges. The thin, transparent walls of both the labrum and the labium are reinforced by chitinous rods. The musculature of the mouth tube consists of two pairs ofretractores oris, four pairs ofcompressores labri and two pairs oflevatores labii. The mandibles are operated by a pair of antagonistic muscles, theroto adductor mandibulae androto abductor mandibulae. Like other lernaeopodids,I. appendiculata resorts to a browsing type of feeding.     

Relating divergence in polychaete musculature to different burrowing behaviors: A study using opheliidae (Annelida)

Divergent morphologies among related species are often correlated with distinct behaviors and habitat uses. Considerable morphological and behavioral differences are found between two major clades within the polychaete family Opheliidae. For instance, Thoracophelia mucronata burrows by peristalsis, whereas Armandia brevis exhibits undulatory burrowing. We investigate the anatomical differences that allow for these distinct burrowing behaviors, then interpret these differences in an evolutionary context using broader phylogenetic (DNA‐based) and morphological analyses of Opheliidae and taxa, such as Scalibregmatidae and Polygordiidae. Histological three‐dimensional‐reconstruction of A. brevis reveals bilateral longitudinal muscle bands as the prominent musculature of the body. Circular muscles are absent; instead oblique muscles act with unilateral contraction of longitudinal muscles to bend the body during undulation. The angle of helical fibers in the cuticle is consistent with the fibers supporting turgidity of the body rather than resisting radial expansion from longitudinal muscle contraction. Circular muscles are present in the anterior of T. mucronata, and they branch away from the body wall to form oblique muscles. Helical fibers in the cuticle are more axially oriented than those in undulatory burrowers, facilitating radial expansion during peristalsis. A transition in musculature accompanies the change in external morphology from the thorax to the abdomen, which has oblique muscles similar to A. brevis. Muscles in the muscular septum, which extends posteriorly to form the injector organ, act in synchrony with the body wall musculature during peristalsis: they contract to push fluid anteriorly and expand the head region following a direct peristaltic wave of the body wall muscles. The septum of A. brevis is much thinner and is presumably used for eversion of a nonmuscular pharynx. Mapping of morphological characters onto the molecular‐based phylogeny shows close links between musculature and behavior, but less correlation with habitat. J. Morphol. 275:548–571, 2014. © 2014 Wiley Periodicals, Inc.     

Structure and anterior regeneration of musculature and nervous system in Cirratulus cf. cirratus (Cirratulidae,Annelida)

Annelids provide suitable models for studying regeneration. By now, comprehensive information is restricted to only a few taxa. For many other annelids, comparative data are scarce or even missing. Here, we describe the regeneration of a member of the Cirratulus cirratus species complex. Using phalloidin‐labeling and antibody‐stainings combined with subsequent confocal laser scanning microscopy, we provide data about the organization of body wall musculature and nervous system of intact specimens, as well as about anteriorly regenerating specimens. Our analyses show that C. cf. cirratus exhibits a prominent longitudinal muscle layer forming a dorsal muscle plate, two ventral muscle strands and a ventral‐median muscle fiber. The circular musculature forms closed rings which are interrupted in the area of parapodia. The nervous system of C. cf. cirratus shows a typical rope‐ladder like arrangement and the circumesophageal connectives exhibit two separate roots leading to the brain. During regeneration, the nervous system redevelops remarkably earlier than the musculature, first constituting a tripartite loop‐like structure which later become the circumesophageal connectives. Regeneration of longitudinal musculature starts with diffuse ingrowth and subsequent structuring into the blastema. In contrast, circular musculature develops independently inside the blastema. Our findings constitute the first analysis of regeneration for a member of the Cirratuliformia on a structural level. Summarizing the regeneration process in C. cf. cirratus, five main phases can be subdivided: 1) wound closure, 2) blastema formation, 3) blastema differentiation, 4) resegmentation, and 5) growth, respectively elongation. Additionally, the described tripartite loop‐like structure of the regenerating nervous system has not been reported for any other annelid taxon. In contrast, the regeneration of circular and longitudinal musculature originating from different groups of cells seems to be a general pattern in annelid regeneration. J. Morphol. 275:1418–1430, 2014. © 2014 Wiley Periodicals, Inc.     

F-actin framework in Spirorbis cf. spirorbis (Annelida: Serpulidae): phalloidin staining investigated and reconstructed by cLSM

Abstract. Serpulidae encompasses polychaete species whose members have fused anterior ends bearing a tentacular crown, a heteronomous segmented body with a thorax and abdomen, and “chaetal inversion” between the two tagmata. The sessile filter‐feeding organisms live in self‐built, coiled, calcareous tubes on algae. The F‐actin muscular subset of Spirorbis cf. spirorbis was stained with phalloidin and three‐dimensionally reconstructed by means of cLSM, aiming to investigate (1) how the tentacular crown is organized and moved, (2) whether the internal structures, e.g., musculature, follow the thorax–abdomen inversion, and (3) whether circular muscles are present in serpulids. The third aim is by reason of recent investigations suggesting that lack of circular muscle fibers may be a common situation rather than a rare variation in polychaetes. In this manner, this article is part of a comparative evaluation of polychaete muscle systems. We found that longitudinal muscles of the body wall project into the tentacular crown, and that radioli and pinnulae possess three muscle types each, facilitating their great mobility. Operculum, collar, and a pair of unidentified organs possess distinct F‐actin filaments. The trunk is mainly moved by five longitudinal muscle strands, most obvious in the abdomen: two dorsal, two ventral, and an unpaired ventromedian one, out of which the dorsal ones are the strongest. In anterior regions, the two dorsal strands form a single continuous layer; the separated strands lessen posteriorly. Solitary transverse fibers are located ventrally in the middle of each segment, stretching between longitudinal muscles and coelomic lining laterally, where they end. Peripheral and central dorsoventral muscles, two pairs per segment each, are present. Circular fibers as well as bracing muscles were not detected. The results indicate that the musculature does not follow the thorax–abdomen inversion and Serpulidae represents the 15th polychaete taxon in which circular fibers are totally missing.     

Functional morphology of the mouth tube of a lernaeopodid Pseudocharopinus narcinae (Pillai, 1962) (Copepoda: Siphonostomatoida)

A detailed account of the structure and musculature of the mouth tube and mode of feeding in P. narcinae is presented. The double-walled mouth tube is formed by a close fitting of the lateral, longitudinal ridges of the labium into the longitudinal groove of the inner, lateral edges of the labrum. The musculature of the mouth tube consists of two pairs of retractores oris, four pairs of compressores labri and two pairs of levatores labii. The host tissues are scraped off by the mandibles repeating a very characteristic movement brought about by roto adductor mandibulae and roto abductor mandibulae pair of muscles. The tissue particles are forced into the buccal cavity with the help of labral muscles. The suction is broken by the contraction of levatores labii group of muscles.     

Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)

The taxonomy of freshwater pulmonates (Hygrophila) has been in a fluid state warranting the search for new morphological criteria that may show congruence with molecular phylogenetic data. We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of Acroloxus lacustris (Acroloxidae), Radix auricularia (Lymnaeidae), and Physella acuta (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of Biomphalaria glabrata consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of Planorbella duryi is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in A. lacustris, P. acuta, and P. duryi, three layers in R. auricularia and Planorbinae and four layers in B. glabrata. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three-layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.     

Unique patterns of longitudinal body-wall musculature in the Acoela (Plathelminthes): the ventral musculature of Convolutriloba longifissura

The ventral musculature of Convolutriloba longifissura (Acoela) has been studied using electron microscopy and fluorescently labeled whole mounts to demonstrate filamentous actin. Attention was directed to the reorganization and renewal of musculature during asexual reproduction and the adaptation of muscle sets for special predatory behavior. Three ventral subepidermal muscle layers could be distinguished in adult C. longifissura: (1) outer circular muscles that encircle the body, (2) intermediate modified longitudinal muscles with concentric pattern around the mouth and V-shaped orientation in the posterior part of the animal, and (3) inner special pore muscles with radial alignment fanning out from the mouth. Additionally, a few very fragile muscles were found at the anterior margin of the animal. The anterior ventral muscle system built a funnel with the mouth opening as organizing center. The special radial muscles and the antagonistically concentric muscles are perfectly adapted to catch prey in such a way that the funnel is put over the prey to press it through the mouth into the digestive syncytium. Convolutriloba longifissura shows a unique way of asexual reproduction by a two-step fission which results in three individuals. Immediately after separation from the mother animal, daughter individuals are missing the concentric and the radial muscle sets around the mouth completely, but within 30 h these sets are renewed for the most part. Two to three days after separation, the mouth opening is visible and the animals move for capturing prey. The peculiar course of longitudinal muscles in C. longifissura with concentric rings anteriorly and a V-shape muscle layer posteriorly shows that the pattern of body-wall musculature in such basal Plathelminthes as the Acoela may be highly modified from the original pattern of longitudinal and circular muscles.