Distribution and prevalence of the predatory planarian Artioposthia triangulata (Dendy) (Trieladida: Terricola) in Scotland

The distribution of the New Zealand flatworm (Artioposthia triangulata) in Scotland was surveyed between July 1991 and February 1993. There were 348 records from domestic gardens, 56 from botanic gardens, garden centres and nurseries, with only 13 from farms. Although most of the records came from around the major cities the flatworm was found to have become established throughout the Scottish mainland and some of the Islands, e.g. Bute, Gigha, Orkney and Skye. The impact of the flatworm on earthworm populations in agricultural land in Scotland was, as yet, found to be minimal but the fact that seven adjacent farms near Dunoon were infected suggested it could be spread from farm to farm and that in the West of Scotland it could become widespread in agricultural land.     

The effect of starvation on size and survival of the terrestrial planarian Artioposthia triangulata (Dendy) (Tricladida: Terricola)

Ten Artioposthia triangulata were kept without food at 5°C, 10°C, 15°C, 20°C and 23°C. Survival and weights were recorded weekly. All planarians died within one week at 23°C and within three weeks at 20°C. One individual survived for 60 wk at 5°C. A negative exponential equation was fitted to average survival times for planarians kept at 5°C, 10°C, 15°C and 20°C. Negative exponential models could also describe the pattern of degrowth by planarians deprived of food. The results are discussed in relation to possible survival of the planarian outdoors in Northern Ireland and the implications for earthworm populations.     

Assessment of the global potential distribution of the predatory land planarian Artioposthia triangulata (Dendy) (Tricladida: Terricola) from ecoclimatic data

Abstract

Artioposthia triangulata was originally described from New Zealand in 1895 but was subsequently found to have spread to Northern Ireland in 1963 and Scotland and England in 1965. It is now widespread in both Ireland and Scotland, where it has been shown to reduce earthworm numbers to below detectable levels. Ecoclimatic data were used in the computer program CLIMEX to estimate the potential spread of A. triangulata to Europe and the rest of the world. Results indicated it could establish in agricultural land in most of north‐western Europe, and persist in domestic gardens throughout much of central Europe, east and west North America, Australia, southern South America, and South Africa. It is difficult to assess either the extent to which earthworm numbers and diversity would be decreased or how far the effect of their loss to soil structure, nutrient cycling, or wildlife would be detrimental in these areas.     

Genetic variation in populations of the terrestrial planarian Artioposthia triangulata (Dendy), and evidence for passive dispersal in Northern Ireland

Genetic variation in 12 populations of the terrestrial planarian Artioposthia triangulata from Northern Ireland and one in the Republic of Ireland were assessed using starch-gel electrophoresis. Seven enzymes - adenylate kinase, aspartate aminotransferase, enolase, malate dehydrogenase, phosphoglucose isomerase, phosphomannose isomerase and superoxide dismutase - provided consistent, resolved stains. Coefficients of genetic identity were similar for all comparisons suggesting that the populations did not belong to distinct taxa. There was no relationship between genetic distance and geographic distance for populations, thus confirming the role of passive dispersal in the spread of A. triangulata. Average percentage polymorphic loci and mean heterozygosities were higher in the east of the Province (41.4% and 0.240), than the west (37.5% and 0.185) suggesting that the planarian had spread westwards after an initial colonisation in the east.     

Studies on Artioposthia triangulata (Dendy) (Tricladida: Terricola), a predator of earthworms

Observations had linked the disappearance of earthworms from a grass field to the presence of Artioposthia triangulata. An experiment demonstrated that this land planarian could severely and quickly reduce numbers of four earthworm species. Two different approaches to sampling A. triangulata were investigated on a grassland site. The first used four different trap types (wood, ceramic tile, 'corriboard' plastic and 5 mm polystyrene beneath a ceramic tile) measuring 15 cm x 15 cm which were compared over a period of 18 wk. The second used one, or two, applications of dilute formalin solution to quadrats followed by counting the residual planarians by hand-sorting the soil beneath the quadrat to a depth of 30 cm. The polystyrene traps were the most effective for detecting the presence of A. triangulata. No planarians were ever found by hand-sorting after two formalin applications and it was concluded that formalin sampling provided a good estimate of the population density. Counts under traps were not related to absolute estimates of population densities so polystyrene type traps should only be used for detection of the planarians. A multiple regression relationship of A. triangulata weight over time in 5% formalin is presented and allows the original liveweight to be calculated from the weight in preservative. Planarian and earthworm population densities in eight fields were sampled and the planarians were found to be randomly distributed. Counts of planarians in 0.25 m² quadrats were not related to numbers of earthworms but overall field population densities were. It is concluded that A. triangulata is a severe threat to the earthworm populations of Northern Ireland.     

Removal trapping to control Artioposthia triangulata

Removal trapping was used to try and reduce numbers of the predatory planarian Artioposthia triangulata in polythene tunnels on a nursery and a grass field. Ten months of trapping in the tunnels failed to have any impact upon flatworm numbers but weekly catches reduced over a period of 12 weeks' trapping in grassland. The calculated residual population in the grassland (40 ha^-1) was small in comparison with that estimated from formalin sampling (940 × 1288 ha^-1). A separate field study examined the effect of trap density on catch by spacing traps at 2.5, 5, 10 and 20 m intervals. This showed that the numbers of planarians caught was inversely related to the logarithm of trap spacing. It is concluded that removal trapping is too demanding of resources to be a viable control option for this pest in commercial and agricultural situations.     

Surface movement rates of the New Zealand flatworm Artioposthia triangulata: potential for spread by active migration

Movement rates for the surface activity of the New Zealand flatworm Artioposthia triangulata (Dendy) (Tricladida: Terricola) were estimated under semi-field conditions in the Faroe Islands. The distance travelled during 10 consecutive 1-minute periods per individual was measured from mucus trails left behind as the flatworms moved over black plastic sheeting sprayed with fine droplets of water. Adult specimens were found to crawl at rates of up to 28 cm min^-1 (almost 17m h^-1), whereas juveniles crawled at rates of between 3 and 4cm min^-1 (approximately 2m h^-1). This study provides the first data concerning the potential for species spread by active migration.     

Ultrastructural observations on rhabdite formation in the planarian, Artioposthia triangulata

The epidermis of the predatory terrestrial flatworm, Artioposthia triangulata has been examined by transmission electron microscopy for the presence of rhabdiform secretions. Two types of secretion are present: epidermal rhabdoids, produced by a special type of epidermal cell and true adenal rhabdites produced by gland cells beneath the epidermis. The epidermal rhabdoids are formed from Golgi-derived vesicles, which fuse together to form the developing rhabdoid. Within the latter is a filamentous network on which granular material is deposited and coalesces to form a rod-shaped inclusion. The rhabdoids accumulate in the apical region of the cell and release their contents from the apical surface. The adenal rhabdites are formed by Golgi-derived vesicles, which become more elongated and their contents more electron-dense as they mature. The vesicles fuse together to form the primordial rhabdite, which continues to lengthen with the addition of further vesicles. The neck of the rhabdite-forming cell passes between the muscle layers and through the basement membrane to open into the base of the epidermal cell. The rhabdites move from the cell body through the neck into the cytoplasm of the epidermal cell and make their way to the apical surface where they are released to the exterior.     

An electron microscope study of primary epidermis formation in freshwater planarian embryos

Early development in freshwater planarians is generally considered to be highly modified to the point of being unique. A careful examination by TEM, however, suggests that the primary epidermis (Skaer, 1965) is formed in a rather regular manner but is partially inverted with respect to the definitive body axes. After formation of the yolk cell syncytium, the blastomeres enclosed within it increase in number in the central area. Some of these blastomeres then move peripherally as a group and fuse to form another syncytium, the primordium of the primary epidermis. This primordium contacts the surface of the yolk-cell syncytium at the place where the primordium will subsequently flow out. The primordium spreads to the opposite pole through the spaces among the syncytial and non-syncytial yolk cell masses.     

The potential spread of terrestrial planarians Artioposthia triangulata and Australoplana sanguinea var. alba to continental Europe

A survey in Great Britain of the introduced terrestrial planarians Artioposthia triangulata and Australoplana sanguinea var. alba, which are obligate predators of earthworms, indicates that after 30 years Artioposthia triangulata is established throughout Scotland. However, apart from an initial record in 1965, Artioposthia triangulata was unrecorded from England until it was found again in 1992; currently there are 25 records.
There are fewer records of Australoplana sanguinea var. alba and these are mainly from the south and west of England. Ecoclimatic data from Edinburgh, where Artioposthia triangulata is common, and Plymouth, close to where Australopiana sanguinea var. alba has been recorded, were used by the computer programme CLIMEX to predict the potential spread of these planarians within Europe. Results suggest that Artioposthia triangulata could become established in north west Europe, including areas of Scandinavia, Germany and Poland, whereas Australoplana sanguinea var. alba would be confined to western Europe, including northern Spain. To slow their spread within Britain and prevent their establishment on continental Europe it is suggested that all nurseries and garden centres which sell containerised plants should initiate and maintain stringent hygiene regimes.     

Structure and maintenance of the epidermis in Friedmaniella sp. (Prolecithophora)

The epidermis of Friedmaniella sp. has been studied using light and electron microscopy. Three main morphological features characterize its cells, namely (1) DNA bodies in the nuclei, (2) an extensive Golgi apparatus with a well-developed system of transport vesicles, (3) clusters of centrioles mainly in the basal cytoplasm and axonemes and rootlets in the middle and apical cell parts. These peculiarities may indicate continuous physiological regeneration within the cell at the level of cell organelles. DNA bodies may prove to be a taxonomic feature distinguishing the Prolecithophora from other turbellarians.     

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.     

Phylogenetic relationships of the Temnocephaloidea (Platyhelminthes)

Temnocephala novaezealandiae (family Temnocephalidae) and Troglocaridicola mrazeki and Scutariella georgica (family Scutariellidae) were studied by electron microscopy in an attempt to reveal characters that would indicate their phylogenetic relationship to other members of the Platyhelminthes. Ultrastructural features of the epidermis in these temnocephaloideans are like those of the neoophoran turbellarians. The epidermis is syncytial, is honeycombed by a multitude of gland necks whose secretions produce an epidermal surface film, and is underlaid by a thick basement membrane. Some cells in the parenchyma are compartmentalized by intrusive cell processes from neighboring parenchymal cells in a fashion similar to parenchymal structure in the Monogenea and Digenea. The spermatozoa have a pair of free 9+1 flagella and contain aligned dense bodies. The Temnocephaloidea is evidently derived from an early rhabdocoel-turbellarian-like ancestor.     

Ultrastructural features of oogenesis in some marine neoophoran turbellarians

An ultrastructural study of oogenesis has been undertaken in some marine species of neoophoran turbellarians belonging to the Tricladida, Proseriata, and Prolecithophora. Among marine triclads, Cercyra hastata has oocytes with a remarkable amount of what appears to be autosynthetic proteinaceous yolk, while Procerodes dohrni and P. lobata have alecithal oocytes. Among the Proseriata, several species of the primitive family Monocelididae (subfamily Monocelidinae) have oocytes with a discrete amount of what appears to be autosynthetic yolk, while Parotoplana macrostyla, of the derived family Otoplanidae, has alecithal eggs. Finally preliminary observations on Plagiostomum maculatum (order Prolecithophora) suggest that oocytes have autosynthetic yolk globules.These results support the hypothesis (previously formulated on the basis of similar ultrastructural investigations on freshwater triclads) that the presence of autosynthetic yolk in some neoophoran turbellarians can be interpreted as a primitive character inherited from an ancestor with archoophoran organization. This plesiomorphic character would still be maintained in some species while lost in others following differentiation of vitellaria which are characteristic of the Neoophora.     

Density of the New Zealand flatworm, a predator of earthworms, in allotments in Edinburgh between 1993 and 1995

The density of the New Zealand flatworm, Artioposthia triangulata, found between 1993 and 1995 on the surface of the soil under various types of debris scattered in four allotments, was 1–2 m^?2. The type of debris did not markedly affect the density of the flatworm and the distribution of the flatworm within the allotment studied was significantly clustered. Fluctuation in numbers of specimens under different sets of compost‐filled plastic sacks varied in a similar manner to one another. The flatworm was most abundant in the upper 100 mm of soil and its egg capsules were most numerous between 100 and 200 mm below the surface. The flatworm was as abundant under the soil surface shelters as it was beneath shelters buried below the soil surface.     

Ultrastructural features of cocoon-shell globules in the vitelline cells of neoophoran platyhelminths

The Neoophora is characterized by the presence of complex female gonads composed of both germaria and vitellaria. The vitellaria are made up of vitelline cells that differentiate to produce and accumulate reserve substances (proteins, lipid, glycogen) and precursors of the egg capsule or cocoon shell (phenolic proteins). A comparative ultrastructural and cytochemical investigation of shell-forming globules from the vitelline cells of some neoophoran platyhelminths shows that the internal structures of the shellforming globules can be grouped into two (or three) main types. In the first, globule content is characterized by a more or less homogeneous electron-dense substructure evidently arising from repeated coalescence of small vesicles produced by the rough endoplasmic reticulum and Golgi complex. A variant of this type of globule shows intermingling (or concentric) electron-dense and translucent areas producing a pattern resembling brain convolutions (convoluted pattern). The second type of shell-globule structure shows a multigranular pattern, presumably resulting from repeated fusions of Golgian vesicles followed by incomplete coalescence of the electron-dense content. Comparison of shell-globule structure in different taxa could be useful in elucidating some complex and still-unclear phylogenetic relationships among Platyhelminthes.     

Mortality of the earthworm Eisenia fetida (Savigny) presented to the terrestrial planarian Ártioposthia triangulata (Dendy) (Tricladida: Terricola)

Ártioposthia triangulata is a terrestrial planarian that feeds on earthworms. Up to 76% of field collected A. triangulata fed on Eisenia fetida earthworms at 10°C in the laboratory. In one experiment, the feeding rate declined from 63% of earthworms attacked to 36% and 34% in the first, second and third feeding opportunities over a week. The average number of E. fetida eaten was 1.4 per planarian per week. The gain in weight of individual A. triangulata was significantly related to the amount of earthworm tissue lost during the feeding process. Over a one week period, the amount of earthworm tissue lost was related to the total amount presented to the planarians but not to their size. Frequency of attack was, however, related to planarian size. The results are discussed in relation to an hypothetical population of earthworms in pasture. It is concluded that the results support the contention that A. triangulata could seriously deplete earthworm populations.     

Studies on the local dispersal of the New Zealand flatworm, a predator of earthworms, in Edinburgh between 1993 and 1999

The flatworm Artioposthia triangulata was found, from studies using weighted down plastic sheeting, to move predominantly through the soil rather than over it and to use earthworm burrows. Under compost‐filled plastic sacks the flatworm was most active at night although its numbers were similar to those during the day. The transfer of specimens into an area covered with weighted plastic sheeting had no lasting effect on their numbers. The flatworm was regularly removed over 6 years from under paving stones, sheets of newspaper and cardboard placed on the ground in a garden. The rise and fall of the numbers of the flatworm under this debris suggest a predator‐prey periodicity between the flatworm and earthworms of 3 years.     

Ultrastructural observations on paracnids. I: Coelogynopora axi Sopott (Turbellaria,Proseriata)

The Schlauchdrüsen or paracnids of Coelogynopora axi Sopott, 1972 consist of two components: a muscle cell and a secretory cell.The secretory cell is provided with a tube, which bears a border of microvilli. In the normal position the tube is situated in the interior of the secretory cell, and the microvilli stand at the inner side of the tube. After expulsion of the tube the microvilli are situated at its free surface.The evagination takes place in response to chemical stimuli and is effected by the contraction of the myofibrils of the muscle cell.The paracnids are supposed to be mechanisms of defense.However, conformities with nematocysts and spirocysts of the cnidarians do not exist.The paracnids in other species of the Coelogynoporidae, for example in Invenusta paracnida (Karling, 1966) and Carenscoilia bidentata Sopott, 1972 differ from those of C. axi in many details.Abbreviations bl- basement lamina - ep- epidermis - hd- hemidesmosomes - mc- muscle cell - mt- microtubules - mv- microvilli - nsc- nucleus of the secretory cell - sb- bowl containing secretion granules - sc- secretory cell - sd- septate desmosome-like structures - sg- secretion granules - t- tube - tf- tonofilaments