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.     

On Pentacoelum hispaniensis,a marine triclad from Spanish freshwater (Platyhelminthes,Tricladida, Maricola)

The marine triclad Pentacoelum hispaniensis is known from a freshwater biotope near Valencia, Spain. Preliminary results suggest a diploid complement of 12 chromosomes for the species. A historical scenario is presented for the occurrence of P. fucoideum in the Caspian Sea and the Baltic Sea, and for the presence of P. hispaniensis in the Mediterranean.     

Towards a phylogenetic classification of dendrocoelid freshwater planarians (Platyhelminthes): a morphological and eclectic approach

We explore and review the taxonomic distribution of morphological features that may be used as supporting apomorphies for the monophyletic status of various taxa in future, more comprehensive phylogenetic analyses of the dendrocoelid freshwater planarians and their close relatives. Characters examined are: (i) musculature of the pharynx; (ii) anatomy of the eye; (iii) distribution of testis follicles along the longitudinal axis of the body; (iv) epidermis of the body wall; (v) musculature of the ventral body wall; (vi) presence or absence of a penis papilla; (vii) position of the mouth opening; and (viii) anterior adhesive organ. A phylogenetic hypothesis, in the form of a character state tree, is developed. This hypothesis suggests that the Kenkiidae should be resurrected as a separate family, sharing a sistergroup relationship with the Dendrocoelidae. This sistergroup relationship is supported by the shared presence of an apomorphic anterior adhesive organ. Although the shape of the anterior adhesive organ, both within and between kenkiids and dendrocoelids, may differ considerably between species, they are all constructed according to the same principles. The monophyly of the Kenkiidae is supported by the presence of an apomorphic thickening of the marginal zone of the epidermis and presumably also by the presence of prepharyngeal testes. Within the Kenkiidae, the genus Macrocotyla should be considered as a junior synonym of Kenkia, which is characterized by the presence of an apomorphic ‘planariid pharynx‐type 2’. The monophyly of the Dendrocoelidae is suggested by the presence of a dendrocoelid type of pharynx. It is shown that multicellular eye cups with numerous photoreceptor cells are not restricted to the Dugesiidae and the Terricola but are also characteristic for the Dendrocoelidae; the phylogenetic implications of this character state distribution are discussed. It is pointed out that in planarians there may be a relationship between: (i) body size, the size of an eye cup and the number of its constituent pigment cells; and (ii) the size of an eye cup and the number of photoreceptor cells that it contains. Some tentative phylogenetic groups are suggested on the basis of the following presumed apomorphic characters: presence of transverse fibres in the ventral subepidermal musculature, presence of an extra longitudinal muscle layer in the outer pharynx musculature, a mouth opening that has shifted anteriad. It is suggested that the presumed monophyly of the genus Bdellocephala is supported by the apomorphic absence of a penis papilla, combined with a penial lumen provided with many plicae.     

The aquatic triclads of the Grozet Islands (Platyhelminthes: Tricladida)

A new species of marine planarian, Obrimoposthia aparala sp. nov. , is described from the Crozet Islands and two other species are newly recorded for these islands, viz. 0. wandeli and Synsiphonium ernesti. The only known freshwater triclad from the Crozets, Dugesia seclusa , is redescribed from the type material. The morphology, ecology, and distribution of these four species is discussed.     

Karyology and karyotype analysis of diploid freshwater planarian populations of the Dugesia gonocephala group (Platyhelminthes,Tricladida) found in Sardinia

Karyology and karyotype analysis were carried out on freshwater planarian populations of the Dugesia gonocephala group. The strains studied were all diploid with chromosomic number 2n = 16; n = 8. They came from 12 sites mainly localized on the west of the island of Sardinia. Three karyotypes indicated with the letters A, B and C were found in which eight homomorphic pairs of chromosomes were easily identified. In karyotype A all chromosomes are metacentric. Ten populations of the twelve examined showed this karyotype which appears to be the most common. In karyotype B the seventh pair of chromosomes is submetacentric. This karyotype is quite common having been previously found in another eight Sardinian localities. Karyotype C differs from the others in having submetacentric third and seventh pairs of the chromosome complement. It was found in only one locality. The differences observed between these three karyotypes could be interpreted either as sign of differentiation at species level, or as an intraspecific variation due to chromosome mutations (pericentric inversions). This revised version was published online in July 2006 with corrections to the Cover Date.     

A new and aberrant species of Dugesia (Platyhelminthes,Tricladida, Dugesiidae) from Madagascar

In this paper we report a new species of Dugesia of the family Dugesiidae from Madagascar, representing the fourth species of freshwater planarian known from this global biodiversity hotspot. In some respects the new species is aberrant, when compared with its congeners, being characterized by a head with smoothly rounded auricles, a peculiar course of the oviducts, including the presence of a common posterior extension, and by the asymmetrical openings of the vasa deferentia at about halfway along the seminal vesicle. Further, it is characterized by a ventral course of the ejaculatory duct with a terminal opening, very long spermiducal vesicles and unstalked cocoons. Its diploid chromosome complement with 18 chromosomes represents an uncommon feature among fissiparous species of Dugesia.     

On the taxonomic status of a Dugesia species from Buenos Aires (Platyhelminthes: Turbellaria: Tricladida)

The taxonomic and distributional statuses of the freshwater planarians of South America are imperfectly understood, and a comprehensive review is sorely needed. The species known from the vicinity of Buenos Aires, Argentina, was first described by Böhmig (1902) under the name Planaria dubia Borelli, 1895. Subsequent authors (Kenk, 1930, 1974; Cazzaniga & Curino, 1987) have cast doubt on this assignation. Some have assigned it to Dugesia anceps Kenk; Hyman (1959) was inclined to regard the Buenos Aires planarian as a geographical variant of Dugesia sanchezi Hyman. A comparison of D. sanchezi from the mountainous region of central Chile with specimens from Buenos Aires confirms that the Argentinian material is a distinct species.The differences between the two species are subtle. The Argentinian specimens are distinctive in their possession of infranucleate epithelia of the atria, penis, and bursal canal, as well as in their penial structure. In both species, the epithelia of the atria and bursal canal are glandular. Those of D. sanchezi are nucleate and their secretions are cyanophilous. In the Argentinian specimens, these same epithelia are infranucleate; but while the secretion of the bursal canal is indeed cyanophilous, that of the atrium is erythrophilous. The presence, in these same specimens, of cyanophilous glands opening into the penial vesicles, and of erythrophilous glands whose secretions contribute to a viscous plug in the ejaculatory duct, suggests a difference in the physiology of copulation.It is apparent that the Argentinian specimens are distinct from D. sanchezi Hyman, and probably they represent a new species, one to be described in detail elsewhere. In any case, Hyman's (1959) suggestion that D. sanchezi occurs near Buenos Aires is mistaken; so also is Cazzaniga's & Curino's (1987) identification of specimens from Buenos Aires as D. anceps.     

Unciliated body surface in three species of the Umagillidae (Dalyellioida,Platyhelminthes)

Anoplodiera voluta Westblad, Seritia elegans (Westblad) and Wahlia macrostylifera Westblad are species of the family Umagillidae living in the intestine of the holothurian Parastichopus tremulus. In all three species, part of the epidermis is unciliated, but unlike unciliated epidermis in the major parasitic flat-worm groups, it is cellular and has intraepithelial nuclei. The surface of the unciliated cells in A. voluta is convex with the cells separated by lateral gaps. The cells have two distinct regions: the basal, organelle-rich part and the apical part which contains few identifiable organelles except vesicles. In W. macrostylifera the unciliated cells have a flat surface and, between them, narrower gaps that in some cases widen to paracellular compartments below the cell surface. Apically the cells contain electron-dense vesicles, often in contact with the surface. S. elegans has unciliated cells separated by gaps. In all these species, apical vesicles indicate secretory activity. Comparison of the epidermis of these three species with the neodermis of the major parasitic flatworm groups within the Neodermata does not support a close relationship of the three to the Neodermata.     

Methods for taxonomic and distributional studies of terrestrial flatworms (Tricladida: Terricola)

Traditionally used methods for collecting and preserving terrestrial flatworms have proved deficient in various respects. Comparison of methods for preparing these animals for taxonomic study has shown that preservation of morphological characters can best be achieved using formaldehyde-calcium-cobalt fixative and subsequent storage in an aqueous solution of propylene phenoxetol plus propylene glycol and that the best-lasting histological preparations make use of light-fast, oxidation-resistant dyes and synthetic mounting media with an anti-oxidant.     

A new species of land planarian from Madeira (Platyhelminthes, Turbellaria, Tricladida)

A new species of terrestrial planarian from Madeira, Kontikia bulbosa sp. nov. , is described. The genus Kontikia is redefined and its scope and status reviewed.     

Repeated DNA elements in planarians of the Dugesia gonocephala group (Platyhelminthes,Tricladida)

The results of the isolation of repetitive DNA elements in the genome of Dugesia etrusca, a species of the Dugesia gonocephala group, are reported. These sequences, about 1.4 kb long, represent only a part of longer interspersed genomic units (De1 family) and appear to be limited to the genome of some planarians of this group, as indicated by a Southern blot analysis performed in different species and populations. The genomic relationships among different species and populations of the D. gonocephala group are discussed in relation to the results obtained in the present work. This revised version was published online in July 2006 with corrections to the Cover Date.     

The karyotypes of Dugesia species from Spain (Turbellaria,Tricladida)

Some species of Planarians, new to Spain, are recorded. Dugesia polychroa, D. sicula, D. iberica and D. gonocephala s. 1. have been investigated karyologically. The former possesses a diploid complement characteristic of the biotype A (2n = 8); the second is diploid with 2n = 18; diploidy and triploidy were found in sexual populations of D. iberica with n = 8. Triploidy occurred in all the asexual strains of the D. gonocephala group with a basic number of either 8 or 9. In this latter case B-chromosomes were occasionally found.     

A phylogeny of the Platyhelminthes: towards a total-evidence solution

We advocate a total-evidence approach for the reconstruction of working phylogenies for the Turbellaria and the phylum Platyhelminthes. Few morphology-based character matrices are available in the systematic literature concerning flatworms, and molecular-based phylogenies are rapidly providing the only means by which we can estimate phylogenies cladistically. Character matrices based on gross morphology and ultrastructure are required and should be internally consistent, i.e. character coding should follow a set of a priori guidelines and character duplication and contradiction is avoided. In order to test our molecular phylogenies we need complementary data sets from morphology. To understand morphological homology we need phylogenetic evidence from independent (e.g. molecular) data. Fully complementary morphological and molecular data sets enable us to validate phylogenetic hypotheses and the combination of these sets in phylogenetic reconstruction utilises all statements of homology. Working phylogenies which include all phylogenetic information not only shed light on individual character evolution, but form a strong basis for comparative studies investigating the origin and evolutionary radiation of the taxonomic group under scrutiny. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular phylogeny of land and freshwater planarians (Tricladida, Platyhelminthes): from freshwater to land and back

The suborder Tricladida (phylum Platyhelminthes) comprises the well-known free-living flatworms, taxonomically grouped into three infraorders according to their ecology: Maricola (marine planarians), Paludicola (freshwater planarians), and Terricola (land planarians). Molecular analyses have demonstrated that the Paludicola are paraphyletic, the Terricola being the sister group of one of the three paludicolan families, the Dugesiidae. However, neither 18S rDNA nor COI based trees have been able to resolve the relationships among species of Terricola and Dugesiidae, particularly the monophyly of Terricola. Here, we present new molecular data including sequences of nuclear genes (18S rDNA, 28S rDNA) and a mitochondrial gene (COI) of a wider sample of dugesiid and terricolan species. The new sequences have been analyzed, together with those previously obtained, in independent and concatenated analyses using maximum likelihood and bayesian methods. The results show that, although some parts of the trees remain poorly resolved, they support a monophyletic origin for Terricola followed by a likely return of some species to freshwater habitats. Relationships within the monophyletic group of Dugesiidae are clearly resolved, and relationships among some terricolan subfamilies are also clearly established and point to the need for a thorough revision of Terricola taxonomy.     

Terrestrial flatworm (Platyhelminthes: Tricladida: Terricola) diversity versus man-induced disturbance in an ombrophilous forest in southern Brazil

Terrestrial flatworms, or Terricola, are sensitive to environmentalchanges and therefore might be excellent indicators of the conservation statusof natural habitats. The present study aimed to answer two main questions: (1)is terrestrial planarian diversity affected by human disturbances, and (2) isthere any species or group of species that indicates such disturbance? The studysite, National Forest of São Francisco de Paula, Brazil, was originallycovered by a mixed ombrophilous forest, but successive reforestations andselective logging have modified the original landscape. We studied Terricoladiversity in the four main habitats in the study area: mixed ombrophilous forest(NA), ombrophilous forest with selective Araucariaangustifolia logging (N), A. angustifoliareforestation (A), and reforestation of Pinus elliottii(P). According to an increasing degree of disturbance, the habitats might beordered as follows: (NA)<(N)<(A)<(P). We conducted 24 surveys in eachhabitat over a period of 1 year. Our results indicate that: (1)Terricola diversity is inversely related to the degree of habitat disturbance;(2) there are species (Geoplana franciscana,Geoplana sp. 5, and possibly Geoplanidae 3 andNotogynaphallia guaiana) that prefer habitats located onthe extreme right along the main axis of a detrended correspondence analysisordination and therefore can be considered as indicators of well preserved,natural habitats. On the other hand there are species(Xerapoa sp. 1, Choeradoplanaiheringi, G. marginata sensu Marcus andGeoplana sp. 2) preferring more disturbed habitats, whichmay form biological indicators of such disturbances.     

A new species of eastmanosteid arthrodire (Pisces: Placodermi) from Gogo, Western Australia

A new species of Eastmanosteus Obruchev is described from the Upper Devonian Gogo Formation of Western Australia as E. calliaspis sp. nov. It is at the pachyosteomorph level of organization having no contact between the plates of the flank and those of the ventral shield behind the pectoral fin. Well preserved material of the parasphenoid and scapulocoracoid throws further light on the structure of these elements in arthrodires. The genus Eastmanosteus is reviewed.