Developmental sequence for the copulatory organs of Kontikia mexicana with remarks on taxonomic significance (Turbellaria: Tricladida: Geoplanidae)

Five specimens, presumably representing different developmental stages of the land planarian Kontikia mexicana (Hyman, 1939), were used to reconstruct the development of the copulatory apparatus in this species. The results support the notion that Kontikia differs from the closely related Caenoplana in its possession of a penis papilla. In the earliest stage available, a penis papilla was absent and other components were not differentiated. In a late-juvenile condition, the gonopore, seminal vesicle, and ejaculatory duct were present. The short penis papilla appeared to arise in this stage by elongation of the terminal tissue around the ejaculatory duct and its separation from the antral wall. The female canal was guarded by an epithelial fold and the glandular duct was present. In the mature condition, the penis papilla was more elongate, and the secretory (prostatic) region of the ejaculatory duct was functional. The female canal, guarded by an epithelial fold, was well-developed with enlarged glandular duct but lacking the posterior diverticulum and the sperm storage system associated with the ovovitelline ducts known in Kontikia orana Froehlich, 1955.     

Selected characters of the copulatory organs in the land planarian family Bipaliidae and their taxonomic significance (Tricladida: Terricola)

Preliminary analysis was made of 76 species in the monotypic family Bipaliidae, for which the copulatory apparatus has been described. Four characters from the copulatory organs were selected: profile of the female organ (three character states), approachment of the ovovitelline ducts to the female organ (two states), shape of the penial papilla (two states), and shape of the male antrum wall (three states). Data were scored for five preliminary ingroup taxa, viz., the restricted genus Placocephalus, and four other a priori defined subgroups within the family, viz., the genus Bipalium sensu stricto and three other informal taxonomic groupings. An artificial outgroup taxon was constructed on the basis of character states generalized from the Geoplanidae subfamilies Caenoplaninae, Pelmatoplaninae and Rhynchodemidae subfamily Microplaninae. Analysis of the data matrix resulted in a single most parsimonious tree with the following topology: (outgroup (Placocephalus (Bipalium, group A (group B1, group B2)))). This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparative ultrastructure of copulatory organs having a stylet in the Proseriata (Turbellaria)

The ultrastructure of male copulatory organs having a stylet has been studied in some genera of the Proseriata.Within the Monocelididae there was a variety of stylet-like hard structures. The stylet in Monocelis fusca was a differentiation of the basement membrane of the epithelium lining a penis-like muscular papilla. The penis papilla in Ectocotyla consisted of circular muscles surrounded by a thickened basement membrane and an epithelium. Archilopsis sp. and Archilina sp. with a duplex copulatory bulb, had a stylet within a spiny cirrus. The stylet in Archilopsis sp. was a cylindrical muscular protrusion with a thickened basement lamina that lined the cirrus lumen. The stylet structure in Archilina sp. was composed of four long spines which were derivatives of the basement membrane. In Ectocotyla multitesticulata and Dupliminona corsicana, the accessory prostatoid organ was provided with a hook-shaped stylet that was differentiated in the basement membrane and of which the material was continuous with the fibrous matrix between the muscles of the prostatic bulb. The stylet and needles in the Archimonocelis species were intracellular differentiations. The copulatory organ in Carenscoilia biforamen consisted of a tubiform stylet and four needles, all of which were also intracellular specializations.I consider copulatory hard structures in the Turbellaria to be taxonomically significant in terms of structure, differentiation, and location (whether subcellular, in the basement membrane, or intracellular).     

A revision of the cosmopolitan land planarian Bipalium kewense Moseley, 1878 (Turbellaria: Tricladida: Terricola)

Knowledge of the cosmopolitan land planarian Bipalium kewense is briefly reviewed. A neotype and paraneotypes are designated. External morphology and internal anatomy of the species are redescribed. Karyological studies on B. kewense reveal a diploid chromosome complement of 16; one subtelocentric, three submetacentric and four metacentric chromosome pairs. Taxonomy of B. kewense is revised and full synonomy provided. New synonomies include Placocephalus isabellinus Geba and Bipalium costaricensis Hyman. Species that phenetically resemble B. kewense are divided into three groups primarily on the basis of stripe pattern and cutaneous musculature. A differential diagnosis distinguishing B. kewense from taxa in these groups is provided. Present distribution of the species is mapped. New records are reported from Barbados, Colombia, Costa-Rica, El Salvador, Indonesia, Madagascar, Malaysia, Mauritius, Mexico, Puerto Rico, Tahiti, Taiwan and Zimbabwe. Worldwide B. kewense exhibits a disjunct distribution pattern. In Australia and in the United States a similar pattern of occurrence is evident. The species appears to be dispersed with rooted plants. The natural range of B. kewense extends from Vietnam to Kampuchea, possibly extending to Malaysia. Elsewhere it has been introduced by man. The bipaliid fauna of Vietnam, to which B. kewense belongs, appears to have closer phenetic affinities with the faunas of adjacent southern rather than northern regions.     

Positional information and gonadal differentiation in the planarian Dugesia lugubris (Turbellaria)

The planarian Dugesia lugubris is a balanced hermaphrodite, meaning that male genetic factors are in equilibrium with female factors. Differentiation of the gonads is controlled by the region in which they develop. According to the classical theory of germ cell formation, these cells stem from neoblasts that are induced to differentiate by factors specific to the gonadal regions, factors presumably due to gradients formed by neurosecretory activity of the cephalic ganglia and longitudinal nerve cords. A more recently proposed theory holds that germ cells in regenerates originate not from neoblasts but from dedifferentiated cells and that characteristics of the gonadal regions are determined by direct interactions of cells here. Results of our experiments with homo- and autoplastic grafst support the classical theory. Prepharyngeal portions grafted onto posterior body portions retained their ability to maintain or induce development of ovaries. Postpharyngeal portions grafted onto anterior portions produced only testes even though the brain developed normally in these regenerates. Under these experimental conditions, gonad regeneration took longer than it does in normal regeneration (i.e., that in which body regions are not displaced).Translated, from the French, by Marianne Klauser and Seth Tyler.     

A new species of land planarian preying on termites in Kenya (Platyhelminthes: Turbellaria: Tricladida: Terricola)

A new species of land planarian is described from Kenya. Microplana rermirophaga sp. nov. preys on termites by capturing them at the openings to their nest. The pharynx is unusually far forwards for a land planarian and this is probably related to the active nature of the prey. There is an anterior depression caused by the termination of parenchymal longitudinal muscle bundles which might act as a sucker. All specimens are immature which leads to some difficulty in assigning the species to a genus.     

Distribution,abundance, and size of rhabdoids in Dugesia polychroa. (Turbellaria: Tricladida)

The ontogenetic changes in the distribution, abundance, and size of rhabdoids were examined in an undisturbed laboratory population of Dugesia polychroa. Irrespective of triclad age, rhabdoids in the epidermis and parenchyma were more abundant on the dorsal than the ventral side of the body. No significant differences were found in the abundance of epidermal or parenchymal rhabdoids among the anterior, medial, and posterior regions of the body. Rhabdoid number and size changed significantly with triclad age, with a marked depression coinciding with the onset of cocoon production. Rhabdoid discharge was correlated with physical and/or physiological disturbance and occurred in the absence of any overt environmental disturbance. Simple allometric relationships were observed between rhabdoid size and number on one hand and body plan area on the other. Different allometric trends were observed from field-collected individuals compared to the undisturbed laboratory population. The potential function of rhabdoids in the Tricladida is discussed in light of these findings.     

Ultrastructural observations on gastrodermal regeneration in the planarian Dugesia japonica (Turbellaria)

The earliest detectable change during regeneration of the gastrodermis in Dugesia japonica was an aggregation of regenerative cells underneath the gastrodermis remaining at the wound margin. The gastrodermal cells in experimental regenerates retained some of their original characters and presented no indication of cell dedifferentiation. The regenerative cells came into contact with the basal surface of gastrodermal cells, forming stratified cell layers. Differentiation of these cells into gastrodermal cells was initiated by the development of synthetic organelles within their cytoplasm. These differentiating cells gave rise to two different types of gastrodermal cells, namely phagocytic cells and sphere cells. In later stages, there was an apparent movement of differentiated gastrodermal cells towards the parenchyma.     

Embryology of the Turbellaria and its phylogenetic significance

Developmental characters — including oocyte and yolk cell structure, patterns of cleavage, and modes of gastrulation — are presented and examined in relation to the phylogeny of the Turbellaria. Eggshell granules, which have been demonstrated to occur in the oocytes of entolecithal eggs and the yolk cells of ectolecithal eggs, are compared among species, and their potential value as a taxonomic character is discussed. The quartet 4d spiral cleavage of the entolecithal egg of polyclads is described as reminiscent of the primitive pattern of early development for the Turbellaria. This is compared to duet spiral cleavage of acoels, and possible phylogenetic schemes involving the two types of spiral cleavage are reviewed. The link between the precise spiral cleavage, which characterizes development of most archoophorans, and blastomere separation (Blastomeren-Anarchie), which occurs in several neoophoran orders, is established by the occurrence of quartet 4d spiral cleavage in one neoophoran order, and of both quartet spiral cleavage and Blastomeren-Anarchie in different species of a second neoophoran order. The epibolic gastrulation of polyclads is described as primitive for the Turbellaria because of its similarity to that of other members of the Spiralia. Although no identical process occurs in neoophoran development, the earlier event of formation of the hull membrane in some neoophorans, and the later event of formation of the definitive epidermis in all neoophorans studied are presented as processes of possible homology to the epibolic gastrulation of polyclads. The lack of correspondence between polyclads and neoophorans in the relationship of the definitive body axes to the egg axis is discussed, and an hypothesis is advanced to account for the differences. The phylogenetic relationships indicated by known developmental phenomena differ only slightly from the scheme presented by Karling in 1974.     

Frontal organs in the Acoelomorpha (Turbellaria): Ultrastructure and phylogenetic significance

Using characters discernible through electron microscopy, we redefine the organ traditionally identified as the frontal organ in acoelomorph turbellarians as being a collection of two to several large mucus-secreting glands whose necks emerge together through a frontal pore at the exact apical pole of the body, i.e. at the point where the pattern of epidermal ciliary rootlets converges. Representatives that we have studied of each of the acoel families Paratomellidae, Diopisthoporidae, Solenofilomorphidae, Convolutidae, Otocelidae, and Mecynostomidae, as well as a representative of the Nemertodermatida, have such glands. Up to five additional types of glands that open anteriorly outside of the frontal pore, some of which are indistinguishable from glands of the general body wall, could be seen in the nemertodermatid, in Hesiolicium inops (Paratomellidae), and in representatives of the latter four acoel families. In Paratomella, three different types of glands open in diffuse fashion in a frontal glandular complex reminiscent of that in the Macrostomida.Sensory elements near the frontal pore appear to be independent of the gland necks, and so the organ cannot be considered a sensory organ.The frontal organ, as described above, appears very likely to be homologous within the Acoelomorpha, and represents another strong (although unrooted) autapomorphy for this line of turbellarian evolution.     

Global diversity of land planarians (Platyhelminthes, Tricladida, Terricola): a new indicator-taxon in biodiversity and conservation studies

Biodiversity conservation requires prioritization of areas for in situ conservation. In that perspective, the present study documents the global diversity of a component of the soil macrofauna, the land planarians, and concerns an exploratory analysis of their possible role as indicators of biodiversity. Diversity is described by three quantitative methods: (1) hotspots of species richness, selecting areas richest in species, (2) hotspots of range-size rarity, identifying areas richest in narrowly endemic species, and (3) complementarity, prioritizing areas according to their greatest combined species richness. The biodiversity measures of species richness and range-size rarity show a great correspondence in the identification of hotspots of diversity; both measures identify the following seven areas as the most biodiverse for land planarians: Sao Paulo, Florianopolis, western Java, Tasmania, Sri Lanka, North Island/New Zealand, and Sydney. It is discussed to what extent the results for the land planarians correspond with those obtained in other studies that assessed biodiversity hotspots for taxa on a global scale. It is noteworthy that land planarians identify a few global hotspots of diversity that generally do not feature, or only have low rankings, in other studies: New Zealand, southeastern Australia, and Tasmania.     

Biology of long slender land planarians (Turbellaria) in Tokyo and environs

The common short-bodied species of Bipalium does not fragment, but individuals of two newly discovered long-bodied species — B. nobile Kawakatsu & Makino, 1982, and B. multilineatum Makino & Shirasawa, 1983 — do regularly fission, usually behind the mouth or genital pore. Some experimental regenerates of these species form rings by adhesion of the anterior with the posterior cut surface. We found two other forms of Bipalium, perhaps representing a further two species, in Hino City, Tokyo, in 1983; and we have preliminarily arranged the forms of Bipalium known in the region into four groups distinguished on the basis of body coloring, position of the mouth, and structure of the copulatory organ.     

Notes on the biology of the freshwater planarian Dugesia bengalensis (Platyhelminthes Turbellaria Tricladida)

Dugesia bengalensis was described by Kawakatsu (Kawakatsu et al., 1983) from specimens collected in West Bengal. We have been studying populations from many different localities in Santiniketan and adjoining areas of West Bengal and can provide additional biological information.The species is hermaphroditic, and its breeding season was found to occur usually between October and March when the winter temperature falls below 25 °C. Outside of the breeding season, D. bengalensis is capable of asexual reproduction by binary fission (Mahapatra et al., 1987).Development of the reproductive organs appeared to be from neoblasts and other mesenchymal cells and, therefore, to be like that typical of other triclads with the exception that some of the neoblasts used for the reproductive tissue appeared to be derived from the intestinal region (Ghosh, 1988; cf. Teshirogi, 1986).During copulation, the partners were oriented in the same direction and not in a head-to-tail position as has been reported (Hyman, 1945) for some planarians.The oval, stalked cocoons were laid in marshy places, and during the period of summer (usually from April to June) they lay dormant in the sandy soil until the onset of the monsoon rains. Then, typically three or four months after they were laid, the cocoons hatched to yield three or four young, a remarkably low number for freshwater triclads (cf Ball & Reynoldson, 1981).     

Genetic divergence in Northamerican freshwater planarians of the Dugesia dorotocephala group (Turbellaria, Tricladida, Paludicola)

The genetic differentiation between the members of the Dugesia (Girardia) dorotocephala group was analyzed by means of multilocus electrophoresis, and comared to that of another planarian secies, D. tahitiensis, also belonging to the subgenus Girardia. The species examined were: D. dorotocephala s.s (2n = 16), D. arizonensis (2n = 8), D. jenkinsae (2n = 8), and the above mentioned D. tahitiensis (2n = 16). The former three species inhabit North America, and show different proportion of fissiparous and sexual individuals; the latter species inhabits Polynesia and is fully asexual. A total of 11 enzyme loci were genetically analyzed: Mdh-1, Mdh-2, Zdh-1, Idh-2, G3pdh, Got-1, Ck, Pgm-2, Ada, Mpi, and Gpi. Low values of observed mean heterozygosity per locus (Ho) were found in the populations studied, ranging from 0 to 0.18 (average 0.08. In asexual populations (except that of D. tahitiensis) fixed heterozygosity was observered in all the individuals for 1 or 2 loci. The genetic divergence between the species examined is very high, with many loci showing discriminating alleles in different taxa (Nei's genetic distance varies from 0.871 to 1.759). The populations of D. dorotocehala s.s., on the contrary, appear to be genetically quite homogenous average D= 0.019), and the genetic distance values are apparently unrelated to their geographic location and to their way of reproduction. The genetic distance between D. tahitiensis, a species not included in the D. dorotocephala group and D. dorotocephala s.s. is 1.314 and hence similar to the D value between two members of;he dorotocephala group: D. dorotocephala and D. jenkinsae (D = 1.303). The genetic relationships among the populations studied were established by UPGMA cluster analysis and multidimensional scaling. The descendence of the North American species with 2n = 8 from a dorotocephala-like ancestor with 2n = 16 is considered. It is suggested that the latter, as well as a tahitiensis-like line, also having 2n = 16, have originated from a common ancestor by geographic isolation.     

Regenerative and reproductive capacities of the fissiparous planarian Dugesia tahitiensis

A pilot study was performed to assess the regenerative capacities of Dugesia tahitiensis Gourbault, 1977, an exclusively fissiparous planarian species. Animals measuring 9.5–12.5 mm in length were used. Head regeneration rate determined by the appearance of eye spots (Brødsted, 1969: 29–46) was extremely high: at 23 °C, it took 43–59 h to regenerate clearly discernible eye spots in 28 specimens. For comparison, 3.8 days were reported for the regeneration of eye spots in 11.9–12.7 mm long D. tigrina (Girard) at 24 °C (Mead, 1985). As all posterior fragments regenerated a head, irrespective of the cutting level, D. tahitiensis seems to match the Phagocata velata (Stinger) type with a head frequency of 100% at every level (Teshirogi et al., 1977; cf. also Brøndsted, 1969:30).     

A histochemical study of the reproductive structures in the flatworm Dugesia leporii (Platyhelminthes, Tricladida)

Abstract. The functional morphology and the topographic distribution of tissues in the reproductive system of specimens of Dugesia leporii , an endemic Sardinian free-living planarian, are investigated. Data are provided on the nature of epithelial and glandular secretions, spermatophores, and cocoons by histochemistry, light microscopy, and scanning electron microscopy. All secreting epithelial cells produce strongly acidic sulfated glycoproteins. Glandular cells secrete strongly acidic sulfated glycoproteins or keratohyalin-like material in the penis bulb, and prekeratin-like material in atrial glands. Secretions of the bursa copulatrix may be involved in the activation of sperm while material produced by the bursa canal and oviducts probably serves to propel spermatophores or sperm and eggs. Mucous secretion of the seminal vesicle may serve to dilute and activate sperm before copulation. The viscous secrete of the ejaculatory duct and vasa deferentia may play a protective role to maintain sperm viability. Materials produced by the penis papilla and atrium probably lubricate the epithelial surface. The bilayered wall of spermatophore made of keratohyalin-like material and strongly acidic sulfated glycoproteins is produced by two gland types of the penis bulb. The bilayered shell of cocoon made of prekeratin-like and keratohyalin-like materials is secreted by both atrial glands and vitelline cells. The cocoon stalk is made of keratohyalin-like material produced by cement glands. Shell glands, producing GAG, are not involved in cocoon formation, but they may be implicated in the dilution and activation of seminal material to favor sperm movement toward the oviducts.     

Organization and differentiation of the body-wall musculature in Macrostomum (Turbellaria,Macrostomidae)

We studied the body-wall musculature, its ECM (extracellular matrix), and the junctional complexes between muscle cells and between muscle cells and ECM in Macrostomum hystricinum marinum Rieger, 1977, using Nomarski-contrast and electron microscopy. Differentiation of these body-wall components was followed by monitoring embryonic stages at 52%, 64%, and 82% of the time between egg-laying and hatching and with study of the hatchling and adult stages. For comparison, the body-wall musculature of other macrostomidans has been examined in conventional light-histological sections.Muscles form a grid of longitudinally, diagonally, and circularly oriented fibers beneath the epidermis in M. hystricinum marinum and this orientation of cells can be found already in embryos at 64% development. Younger embryos at 52% development show no muscle differentiation. The ECM forms a net-like arrangement that apparently envelops the individual muscle cells. Characteristic knob-like thickenings of the ECM occur at the base of the epidermis. Muscle cells attach to each other, to the epidermis, and to other cell types through hemidesmosome-like junctions at thickenings of the ECM in the adult and hatchling stages; no true desmosomes exist between muscle cells. Gap junctions occur commonly between longitudinal muscles of adult specimens and between perikarya of muscle cells in embryos at 64% and 82% development.More comparative studies are needed to determine the systematic value of presence or absence of the diagonal muscle fibers in the body wall of turbellarians.     

Morphology and histology of male and female reproductive systems in the inseminating species Scoloplax distolothrix (Ostariophysi: Siluriformes: Scoloplacidae)

The morphology and histology of male and female reproductive systems were examined in Scoloplax distolothrix. Internal insemination was documented in this species by the presence of sperm within the ovaries. Mature males and females have elongated genital papillae, exhibiting a tubular shape in males and a plain heart‐shape with two median protuberances in females. The testes are two elongated structures that converge ventrally, under the intestine, towards the genital papilla. They are joined at the caudal end, forming an ovoid single chamber for sperm storage. Secretory regions were not observed. In the lumen of the testicular tubules, spermatozoa can be tightly packed along their lengths, but do not constitute a spermatozeugmata. The lumen of the sperm storage chamber and spermatic duct are filled with free spermatozoa without the accompanying secretions. The ovaries are bird‐wing shaped, saccular structures that converge ventrally under the intestine, towards the genital papilla. They are joined at the caudal end, forming a tubular chamber possibly destined for oocyte storage. An oviduct with an irregular outline connects the chamber to the tubular region of the genital papilla. No distinct sperm storage structure was found in the ovaries. The unique male and female genital papillae suggest that these structures are associated with the reproductive mode in scoloplacids, representing evidence for insemination. The occurrence of free spermatozoa, without the accompanying secretions and not arranged in a spermatozeugmata can be associated with the presence of a tubular male genital papilla for sperm transfer to the female genital tract. This reinforces the idea that sperm packets are not necessary for all inseminating species. The male reproductive system in scoloplacids is very different from that in auchenipterids, a second catfish family with insemination, which indicates that the occurrence of insemination is not connected to the internal morphology of reproductive organs. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Records of the land planarian Polycladus gayi (Tricladida,Geoplanidae) preying on black snails Macrocyclis peruvianus (Gastropoda,Macrocyclidae)

Polycladus gayi is a colourful and easily recognizable land planarian native to the temperate to cold rainforests of the west coast of southern South America, in Chile and Argentina; however, its ecology is poorly understood. Here, we describe the first records of P. gayi preying on a black snail Macrocyclis peruvianus in the wild. Two events were recorded, the first in a healthy forest in Llancahue Alto, Panguipulli, Chile, during the austral summer of 2023, and the second in a mature evergreen forest in the coastal range of La Union, Chile. These records expand our knowledge of the feeding habits and behaviour of P. gayi and add M. peruvianus to its prey list.