A new species of giant planarian from Lake Baikal,with some remarks on character states in the Dendrocoelidae (Platyhelminthes,Tricladida, Paludicola)

On the basis of newly collected material the subspecies Bdellocephala angarensis bathyalis Timoshkin & Porfirjeva, 1989 is raised to full species status, B. bathyalis Timoshkin & Porfirjeva, 1989. Specimens of this species have been collected in Lake Baikal from depths ranging between 610 and 1060 m. The species is characterized by a light, uniform brown pigmentation, absence of eyes, distinct atrial folds, and large size. It is suggested that two features may be useful in elucidating the phylogenetic relationships between dendrocoelid genera: pharynx musculature, and presence of an extra layer of circular muscle in the ventral subepidermal body musculature. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phylogeny of the family Congiopodidae (Perciformes: Scorpaenoidea), with a proposal of new classification

The phylogenetic relationships of the family Congiopodidae are inferred based on morphological characters. The monophyly of this family is supported by 13 unambiguous apomorphic characters, including four autapomorphies among the superfamily Scorpaenoidea. The Congiopodidae shares 26 apomorphic characters with other scorpaenoid taxa, and these characters are considered to also support the monophyly of the family. Upon completion of the phylogenetic analysis using the characters in 39 transformation series, it was assumed that the family is unambiguously supported by five characters (and also by three and one characters when ACCTRAN and DELTRAN are used, respectively) and is branched into two major clades, including Congiopodus and Alertichthys plus Zanclorhynchus, respectively. Based on the phylogenetic relationships, a new classification, recognizing two subfamilies (Congiopodinae and Zanclorhynchinae) in the family Congiopodidae, is proposed. The genus Perryena, that was recently inferred being closely related to the Tetrarogidae (although many authors included it in the Congiopodidae), is provisionally placed into the Congiopodidae as incertae sedis.     

The limits and relationships of the acanthomorph teleosts

The phylogenetic status of the speciose euteleostean assemblage, the Acanthomorpha, is investigated. For the purposes of cladistic analysis attention is centred upon an examination of euteleostean buccal ligament systems, the occipital and ethmoid regions of the neurocranium, and the posttemporal/neurocranial association.
In this study four derived morphological features are identified which support the monophyly of the Acanthomorpha, and the previously 'problematical' genus Polymixia is determined to be the sistergroup of all the remaining acanthomorphs. The phylogenetic integrity of the sistergroup of the entire Acanthomorpha, the Myctophiformes (sensu Rosen, 1973), is attested by the presence of four derived features, and the monophyly of its major subdivisions (the Myctophidae and Neoscopelidae) is resolved.
Additional problems of euteleostean systematics and systematic methodology are discussed.     

Ultrastructure and function of the pharynx of Gnathostomula paradoxa (Gnathostomulida)

 The pharynx of Gnathostomula paradoxa consists of the partly syncytial pharyngeal musculature, a pharyngeal epithelium, myoepitheliocytes, receptors, nerves, and three solid parts, called the jugum, the basal plate, and the jaw. Extended non-contractile regions of both pharyngeal and body wall musculature form the so-called parenchymatous tissue between the digestive tract and the body wall. The pharyngeal epithelium mediates the force from the pharyngeal musculature to the solid parts. The basal plate and jaw contain longitudinal cuticular rods which are elastic antagonists of the musculature. There is no buccal ganglion in G. paradoxa. The study supports the monophyly of the Gnathostomulida and Gnathifera. Accepted: 4 April 1997     

The neuroactive substances and associated muscle system in Rhipidocotyle campanula (Digenea,Bucephalidae) from the intestine of the pike Esox lucius

We report about the muscular system and the serotonergic and FMRFamidergic components of the nervous system of the Bucephalidae trematode, Rhipidocotyle campanula, an intestinal parasite of the pike. We use immunocytochemical methods and confocal scanning laser microscopy (CLSM). The musculature is identified by histochemical staining with fluorescently labeled phalloidin. The body wall musculature of R. campanula contains three layers of muscle fibres – the outer thin circular, intermediate longitudinal and inner diagonal muscle fibres running in two opposite directions. The digestive system of R. campanula possess of a well-developed musculature: radial, longitudinal and circular muscle elements are detected in the pharynx, circular and longitudinal muscle filaments seen in the oesophagus, and longitudinal and the circular muscle fibres were found in the intestinal wall. Specific staining indicating the presence of actin muscle filaments occurs in the cirrus sac localized in the posterior body region. The frontal region of anterior attachment organ, the rhynchus, in R. campanula is represented by radial muscle fibres. The posterior part of the rhynchus comprise of radial muscles forming the organ's wall, and several strong longitudinal muscle bundles. Serotonergic and FMRFamidergic structures are detected in the central and peripheral compartments of the nervous system of R. campanula, that is, in the paired brain ganglia, the brain commissure, the longitudinal nerve cords, and connective nerve commissures. The innervations of the rhynchus, pharynx, oesophagus and distal regions of the reproductive system by the serotonergic and FMRFamidergic nervous elements are revealed. We compare our findings obtained on R. campanula with related data for other trematodes.     

Rhipidocotyle genovi n. sp. (Digenea: Bucephalidae) from the intestine ofGaidropsarus mediterraneus (L.) (Gadiformes: Gadidae) from the Black Sea

Rhipidocotyle genovi n. sp. is described from the fishGaidropsarus mediterraneus (L.) (Gadiformes: Gadidae) in the Black Sea. Morphological features characteristic of this species include: an anterior sucker with seven prominent muscular appendages; a pharynx rather posterior in position; ovary and testes relatively anterior in position, usually anterior to the pharynx; a long (in relation to body length) sucker, cirrus-sac and excretory vesicle; the presence of a pre-oral lip; and contiguous vitelline fields which form an arch. The new species is distinguished fromR. galeata, R. lamberti andBucephalus marinum. This is the first member of the genus whose adults are recovered from Black Sea marine fishes. Metacercariae, which encyst in large numbers in the musculature and fins of the blenniid fishesBlennius tentacularis, B. sanguinolentus, B pavo, B. sphinx andB. zvonimiri and exhibit a similar morphology to the adult, are described.     

Predation behaviour of land planarians

Predatory behaviour of land planarians is seldom observed or reported. Aspects reported are (1) finding prey; (2) attack behaviour; (3) capture using adhesive mucus, pharyngeal action, poisonous secretions, physical embrace; (4) feeding by extension of pharynx, releasing copious digestive fluid. The species Bipalium kewense, B. adventitium and B. pennsylvanicum attack earthworms, immobilizing them by physical holding, digesting by pharyngeal secretions and then ingesting the treated tissue. Group attacks on giant African land snails involving chemotactic tracking, occur in Platydemus manokwari and Endeavouria septemlineata. Specialized capture methods are used by some species; Rhynchodemus sylvaticus uses an expanded cephalic hood to capture small insects and in Africa, termites are captured by the elongated anterior of Microplana termitophaga as planarians wait within the colony air shaft openings to ensnare the workers in sticky mucus. The result of extensive predation by land planarians may seriously reduce the prey, e.g., providing effective population control of giant land snails by introduced Platydemus manokwari, or causing serious depletion of desirable earthworm populations by the exotic Artioposthia triangulata in North Ireland.     

The adult musculature of two pseudostomid species reveals unique patterns for flatworms (Platyhelminthes,Prolecithophora)

We analyzed the adult musculature of two prolecithophoran species, Cylindrostoma monotrochum (von Graff, 1882) and Monoophorum striatum (von Graff, 1878) using a phalloidin-rhodamine technique. As in all rhabdithophoran flatworms, the body-wall musculature consisted of three muscle layers: on the outer side was a layer of circular muscle fibers and on the inner side was a layer of longitudinal muscle fibers; between them were two different types of diagonally orientated fibers, which is unusual for flatworms. The musculature of the pharynx consisted of a basket-shaped grid of thin longitudinal and circular fibers. Thick anchoring muscle fibers forming a petal-like shape connected the proximal parts of the pharynx with the body-wall musculature. Male genital organs consisted of paired seminal vesicles, a granular vesicle, and an invaginated penis. Peculiar ring-shaped muscles were only found in M. striatum, predominantly in the anterior body part. In the same species, seminal vesicles and penis only had circular musculature, while in C. monotrochum also longitudinal musculature was found in these organs. Female genital organs were only present in M. striatum, where we characterized a vagina interna, and a bursa seminalis. Transverse, crossover, and dorsoventral muscle fibers were lacking in the middle of the body and greatly varied in number and position in both species.     

Muscles of the pes of hylid frogs

Complete or partial dissection of the foot musculature of 404 hylid frogs representing 247 species and 33 genera, along with representatives of eight other families, revealed a number of apomorphic characters that distinguish the hyloid frogs (Hylidae plus Allophryne and Centrolenidae) from other bufonoid frogs. Additional characters were found to define some of the hylid subfamilies. Addition of characters from the foot musculature to Duellman's phylogenetic tree of the hyloids produced a tree in which Allophryne and Centrolenidae are nested within Hylidae. Support was found for the monophyly of the 30-chromosome group within Hyla, and for a large number of the groups that comprise "Boana," viz., the Hyla albomarginata, H. albopunctata, H. boans (except H. vasta), H. geographica, and H. pulchella groups, but foot muscle characters provide no information relating to relationships of the West Indies hylines.     

Musculature of the facultative parasite Urastoma cyprinae (Platyhelminthes)

In an effort to understand how the feeding motions of Urastoma cyprinae are generated, the arrangement of its musculature was studied using fluorescence microscopy of phalloidin‐linked fluorescent stains and conventional light histology and transmission electron microscopy. BODIPY 558/568 phalloidin and Alexa 488 phalloidin resolved a meshwork of ribbon‐shaped body‐wall muscles as well as inner‐body musculature associated with the pharynx and male copulatory organ. The general pattern of body‐wall muscles in U. cyprinae is similar to that of other rhabdocoel turbellarians in consisting only of circular, longitudinal, and diagonal fibers; the arrangement of these muscles readily correlates with the bending motions the animal undergoes as it feeds at the surface of gills in bivalves it parasitizes. The orogenital atrium of U. cyprinae lies at the posterior apex of the body, opening at a terminal pore. As evidenced by the arrangement of its epithelium and musculature, it appears to be an invagination of the body wall and comes closest of any such duct studied in turbellarians to satisfying the hypothetical model of a “pseudopharynx,” ostensibly adapted as an organ for swallowing and so supplementing the ingestive role of the animal's true pharynx. J. Morphol. 241:207–216, 1999. © 1999 Wiley‐Liss, Inc.     

Molecular analysis of the diversity of terrestrial planarians (Platyhelminthes, Tricladida, Continenticola) in the Iberian Peninsula

This work is a prospective study to estimate the potential species diversity of terrestrial planarians in the Iberian Peninsula. Live specimens were collected from several Iberian localities and assigned to different morphotypes on the basis of their external morphological characteristics. From the same specimens, sequences from the mitochondrial Cytochrome Oxidasa subunit I (COI) and from the nuclear ribosomal gene 18S rRNA were obtained. Sequences from GenBank of the families Dendrocoelidae, Planariidae, Bipaliidae, Geoplanidae and Rhynchodemidae have been used as outgroups in the phylogenetic analysis. The results showed that terrestrial planarians have a wide distribution in the Peninsula, with all individuals found belonging to the Rhynchodemidae family. Morphological observations indicated the presence of 10 morphospecies confirmed by the molecular analyses. At the same time, COI sequences were successfully used as a molecular marker for species identification in the barcoding mode, which is of great use in groups like this with few external morphological characteristics. The combined data strongly suggest the presence of at least 15 species in the Iberian Peninsula, a number that nearly doubles previous estimates, indicating that terrestrial planarians are more diverse than expected in the region and, as proposed, may be a good biodiversity indicator and model for biogeographical studies.     

The triradiate sucking pharynx in animal phylogeny

Independent sucking/swallowing pharynges, as found in members of at least ten animal phyla, are pharynges that are opened by muscles that are not attached to the outer body wall. Their musculature is derived from either mesoderm or ectoderm, the latter taking the form of a myoepithelium. I review results of previous work on the morphology of independent sucking/swallowing pharynges among invertebrates and provide new information on the ultrastructure of the pharynx of the cyclostome bryozoan Crisia eburnea. The various morphologies of this type of pharynx have been used in some phylogenetic considerations, but only the myoepithelial sucking pharynx with a triradiate lumen was considered significant in analyses of relationships between phyla. However, I argue that this shape is the only one that makes an efficient suction pump, and this, together with different orientations of the myoepithelial pharyngeal pump and its phylogenetic distribution, indicates that the triradiate myoepithelial pharynx has evolved convergently in a number of lineages.     

Revision of the Childiidae (Acoela), a total evidence approach in reconstructing the phylogeny of acoels with reversed muscle layers

The Childiidae sensu Dörjes 1968 comprises the acoel worms characterized by a cone‐shaped penis with muscular or sclerotized elements. Based on differences in body‐wall musculature arrangement, Hooge (2001) recently restricted the family to the genus Childia Graff, 1910 and placed the remaining genera to his new family Actinoposthiidae Hooge 2001 . This rearrangement has been questioned ( Raikova et al. 2004 ). We reconstructed the phylogeny of the Childiidae sensu Dörjes 1968 by means of a total evidence analysis including Histone H3, 28S rDNA and new 18S rDNA sequences, as well as 50 morphological characters. New characters of the muscular system and copulatory organs discovered through confocal laser scanning microscopy of phalloidin‐stained specimens are included in the phylogenetic analysis. A total of 12 taxa (nine ingroup and three outgroup) were used in the parsimony analysis of the 18S data set, which was aligned with different parameters for a sensitivity analysis, and the combined data set (18S + 28S + H3 + morphology). Incongruence in the node support of the groups among the four partitions was very low in the total evidence tree; except for the H3 partition. The conflict observed in the H3 partition is likely due to large homoplasy observed in the synonymous alternatives at both first and third codon positions. All data partitions demonstrated that Actinoposthia beklemischevi Mamkaev 1965 , and the newly defined taxon Childiidae (comprising Childia and Paraphanostoma Westblad 1942 ) are not close relatives. The monophyly of Childia and Paraphanostoma is strongly supported by both the 18S and 28S data partitions. Our study also reveals additional apomorphies uniting Childia with Paraphanostoma from body‐wall musculature, statocyst muscles and male copulatory organ. Muscular system, statocyst muscles, male copulatory organ and nervous system characters proved to be the best characters for taxonomic delimitations of subtaxa within the Childiidae, whereas the seminal bursa (a frequently used character in the taxonomy of Acoela) was highly homoplastic. We also described the body‐wall musculature of six Paraphanostoma species, which is characterized by the reversed arrangement of the longitudinal and circular muscle layers, and by the absence of diagonal muscles on the ventral side of the body and the presence of two types of diagonal muscles on the dorsal side. Childia groenlandica (Levinsen, 1879) is nested among the Paraphanostoma species in our total evidence tree, so we synonymize Paraphanostoma with Childia; all former members of Paraphanostoma are transferred to Childia.     

Two new species of creagrocercid nematodes parasitic in earthworms,with comments on the phylogenetic affiliations of the Creagrocercidae Baylis, 1943

Two new species of the rare nematode family Creagrocercidae from earthworms are described and illustrated. Creagrocercus braziliensis n. sp. is distinguished from the type-species, C. barbatus Baylis, 1943, by: the presence of four cephalic papillae (vs a pair of finger-shaped latero-ventral processes) on the head; larger amphids; a more posterior excretory pore position; a shorter pharynx which is remarkably expanded at the base; the nerve-ring situated just posterior to the base of the pharynx (vs at mid-pharyngeal level); an anal aperture present in the females; anterior ovary and testis reach just into the anterior half of the body (vs closely posterior to the pharynx); equal, similar (vs unequal, dissimilar) spicules; and a prominent, unpaired precloacal papilla in males. Creagrocercus drawidae n. sp. is related to C. barbatus by a similar pharyngeal shape and the presence of a tail ‘hook’, and to C. braziliensis n. sp. by: the lack of finger-shaped processes on the head and the presence of four cephalic papillae; a similar position of the excretory pore; a similar arrangement and length of the reproductive system; the presence of unpaired precloacal papilla in males; similar, almost equal spicules; and the presence of a tail ‘hook’. From both of these species C. drawidae differs by having: a much shorter body and pharynx; larger amphids; a more posterior nerve-ring position; larger and less numerous eggs; greatly inflated (vs flat) vulval lips; shorter spicules; and a smaller caudal ‘hook’. For C. drawidae, the partial sequences of the SSU (18S) rDNA and the D2D3 segment of LSU (28S) rDNA were obtained and subjected to phylogenetic analyses. The phylogenetic affiliations of the Creagrocercidae are discussed.     

A molecular analysis of the phylogenetic position of the suborder Cavernicola within the Tricladida (Platyhelminthes), with the description of a new species of stygobiont flatworm from Benin

This paper presents a new species of cavernicolan planarian from the African continent, which enabled us to determine the position of the suborder Cavernicola within the order Tricladida on a molecular basis. A recent paper suggested a sister‐group relationship between Cavernicola and marine triclads. However, our work is the first molecular study to demonstrate unequivocally that the Cavernicola does not group with either the freshwater planarians or the land planarians, i.e. is not closely related to the suborder Continenticola, and corroborates the suggested sister‐group relationship with the marine triclads or Maricola. Two possible scenarios are proposed for the evolution of the Cavernicola from marine ancestors. The new species is assigned to the genus Novomitchellia and is characterized by the absence of eyes. Testicular follicles are numerous and only dorsal, extending from behind the ovaries to the posterior end of the body. The vasa deferentia separately penetrate the penis bulb and the seminal vesicle. The penis papilla is very short and blunt, and the penis bulb musculature is weak. A long and narrow copulatory bursa is situated behind the gonopore. The common oviduct is orientated perpendicularly to the short bursal canal.     

The ultrastructure of the marine gastrotrichTurbanella cornuta Remane (Macrodasyoidea) and its functional and phylogenetical importance

Summary The organization of marine gastrotrichs (Macrodasyoidea) is reviewed by ultrastructural analysis of one representative,Turbanella cornuta Remane, and the fine structure of tissues and cells is described. Turbanella cornuta has a mono-layeredcellular epidermis rich withsensory hairs, epidermal bodies, isolatedepidermal glands, glandular adhesive organs belonging to a duo-gland type, andventral ciliated epidermal cells of the multiciliated type. The voluminous neuropil of thebrain consists of a circular commissure which sends out four anterior and posterior longitudinal headnerves. The posterior ones unite on each side to one single longitudinal nerve of the periphery which is occupied with single peripheral neurons and has thin commissures that make it anorthogon. The position and the structure of the neurons indicate their sensitive, associative, motoric, and neurosecretory functions. The different forms of synapses give first hints to neuronal connections within gastrotrichs. There is a big cellularglia around the brain commissure and a small cellular glia within the brain neurons. In between the cross-striated muscle fibrils of thepharyngeal wall there are also nerves and sensory hairs.TheY-organ lies in the interior of the lateral body cavities, which are delimited by an outer musculature of the body wall and an inner musculature of the intestinal tract. In the pharyngeal region, theY-organ fills the body cavities completely and, in the intestinal region, it covers thegonads, which also lie in the lateral body cavities, dorsally. The testicles lie separately in front of the paired ovaries. Single states of oogenesis could be identified as oogonia, and young and old oocytes. There is a paired gland organ in front of the dorsomedian ovary which may produce a mucous cover for the egg.Theintestinal tract is adapted to mechanical stress by a myoepithelium in the pharyngeal region, by various interdigitations, and by narrow intercellular gaps with hemidesmosomal adhesions to the basement membrane. The majority of the resorbing intestinal cells have a high seam of microvilli and contain various numbers of lysosomes. In addition, there are some secerning cells without microvilli, but with a centrically arranged ER and with big secretion granules in the dorsomedian sector.The ultrastructure affirms a close correlation between the conditions of life in the interstitium and structural adaptations, such as may be observed in single structures of the body wall, the y-organ, the intestinal tract and, in some respect, even in the nervous system and in the formerly researched musculature and spermatohistogenesis. On the other hand, for the construction of the glandular adhesive organs, the nervous system, and the formerly investigated body cavities, a phylogenetical relevance is discussed. Thereafter, gastrotrichs have more primitive characters than the closely related nematodes.Abbreviations a sensory hair cells - am ampoule - at outleading tube - b basement membrane - bb basal body - c cilium - cr rootlet of the cilium - cu cuticle - cw cell wall - d d-cells of the brain - de desmosomes - e e-cells of the brain - eb epidermal bodies - ee ripe egg in the dorsomedian ovary - ep epidermis - er endoplasmatic reticulum - ev ventral ciliated epidermal cells - f f-cells of the brain - fr fibrillar structure - g gland cell - ge germ epithelium - gl₍₁₊₂₎ small and big cellular glia of the br - go Golgi-apparatus - gp genital pore - h h-cells of the brain - hf lateral adhesive tubules - hfp posterior adhesive tubules - i intestine - il intestinal lumen - 1 lumen of the organ - li lipid granules - ly lysosomes - m mitochondrium - mb multivesicular body - mc circular musculature - mi microvilli - ml longitudinal musculature - mo mouth opening - mt microtubules - mpl longitudinal muscle fibers of the pharyngeal wall - mpr radial muscle fibers of the pharyngeal wall - n nucleus - nb brain neurons - nc brain commissure - nf nerve fibers - nl lateral headnerve - nm nuclear membrane - nn nucleolus - nv ventrolateral headnerve - nz peripheric neuron - ncp peripheric nerve commissure - nvp longitudinal peripheric nerve - o lateral ovary - oc oocyte - oo oogonium - ow wall cells of the ovary - p secretory pore - ph pharynx - po palpar organ - phb pharyngeal bulbs - phl pharyngeal lumen - phn nerve plexus of the pharynx wall - sa anterior sense organ - sg secretory granules - sh sensory hair cell - sp posterior sense organ - st supporting stick - su supporting cell - sv synaptic vesicles - sy synaptic gap - t testicles - tl testicular lumen - tw wall cells of the testicles and the vas deferens - v ventral - va vacuoles - vd vas deferens - vs vesicles - y y-organ - yc anterior commissure of the y-organ - z yolk granules     

Quasimodoana, a new Holarctic genus of eucoiline wasps (Hymenoptera, Cynipoidea, Figitidae), with a phylogenetic analysis of related genera

Abstract.  A new Holarctic genus, Quasimodoana gen.n. is described for the Palearctic eucoiline wasp Eucoila decipiens Förster, 1869, as Quasimodoana decipiens comb.n. A lectotype is designated for this species. A new North American species is described here as Quasimodoana gibba sp.n. As the new genus has similarities with several eucoiline genera, belonging to two related but distinct lineages, a phylogenetic analysis was carried out, based on 83 morphological characters. The two lineages included in the analysis are referred to here under the tribal names of Eucoilini and Trichoplastini stat.nov. We discuss reasons for adopting available tribal names (rather than 'informal genus groups'), and Trichoplastini is removed from synonymy with Eucoilini. The phylogenetic analysis places Quasimodoana unambiguously within Eucoilini, as a sistergroup to the Trybliographa complex. However, there is only weak support for the monophyly of Eucoilini and for the hypothesized sistergroup relationship between Eucoilini and Trichoplastini.     

A robust molecular phylogeny of the Tricladida (Platyhelminthes: Seriata) with a discussion on morphological synapomorphies.

The suborder Tricladida (Platyhelminthes: Turbellaria, Seriata) comprises most well-known species of free-living flatworms. Four infraorders are recognized: (i) the Maricola (marine planarians); (ii) the Cavernicola (a group of primarily cavernicolan planarians); (iii) the Paludicola (freshwater planarians); and (iv) the Terricola (land planarians). The phylogenetic relationships among these infraorders have been analysed using morphological characters, but they remain uncertain. Here we analyse the phylogeny and classification of the Tricladida, with additional, independent, molecular data from complete sequences of 18S rDNA and 18S rRNA. We use maximum parsimony and neighbour-joining methods and the characterization of a unique gene duplication event involving the Terricola and the dugesiids to reconstruct the phylogeny. The results show that the Maricola is monophyletic and is the primitive sister group to the rest of the Tricladida (the Paludicola plus the Terricola). The Paludicola are paraphyletic since the Terricola and one paludicolan family, the Dugesiidae, share a more recent common ancestor than the dugesiids with other paludicolans (dendrocoelids and planariids). A reassessment of morphological evidence may confirm the apparent redundancy of the existing infraorders Paludicola and Terricola. In the meantime, we suggest replacing the Paludicola and Terricola with a new clade, the Continenticola, which comprises the families Dugesiidae, Planariidae, Dendrocoelidae and the Terricola.