A new subfamily, new genus and new species of terrestrial flatworm (Platyhelminthes: Tricladida: Geoplanidae) from Stewart Island, New Zealand

Within the context of ongoing taxonomic revisions of terrestrial flatworms from the Australia–New Zealand region: the new genus Eudoxiatopoplana is erected; the new species Eudoxiatopoplana bilaticlavia is described from Stewart Island; and the new subfamily Eudoxiatopoplaninae is erected within the family Geoplanidae

Intracellular immunohistochemical detection of tetrodotoxin in Pleurobranchaea maculata (Gastropoda) and Stylochoplana sp. (Turbellaria)

Tetrodotoxin (TTX), is a potent neurotoxin targeting sodium channels that has been identified in multiple marine and terrestrial organisms. It was recently detected in the Opisthobranch Pleurobranchaea maculata and a Platyhelminthes Stylochoplana sp. from New Zealand. Knowledge on the distribution of TTX within these organisms is important to assist in elucidating the origin and ecological role of this toxin. Intracellular micro-distribution of TTX was investigated using a monoclonal antibody-based immunoenzymatic technique. Tetrodotoxin was strongly localized in neutral mucin cells and the basement membrane of the mantle, the oocytes and follicles of the gonad tissue, and in the digestive tissue of P. maculata. The ova and pharynx were the only two structures to contain TTX in Stylochoplana sp. Using liquid chromatography-mass spectrometry, TTX was identified in the larvae and eggs, but not the gelatinous egg cases of P. maculata. Tetrodotoxin was present in egg masses of Stylochoplana sp. These data suggest that TTX has a defensive function in adult P. maculata, who then invest this in their progeny for protection. Localization in the digestive tissue of P. maculata potentially indicates a dietary source of TTX. Stylochoplana sp. may use TTX in prey capture and for the protection of offspring

First report of the land planarian Endeavouria septemlineata (Hyman, 1939) (Platyhelminthes, Tricladida, Continenticola, Geoplanidae) in French Polynesia.

We report the presence of the land planarian Endeavouria septemlineata from Tahiti, French Polynesia, on the basis of a single specimen collected in 2017. Identification of the species was ascertained by external and internal morphology and DNA COI sequence. The finding is of importance for conservation, since this species is a predator of soil animals

Giant worms chez moi! Hammerhead flatworms (Platyhelminthes, Geoplanidae, Bipalium spp., Diversibipalium spp.) in metropolitan France and overseas French territories

Background: Species of the genera Bipalium and Diversibipalium, or bipaliines, are giants among land planarians (family Geoplanidae), reaching length of 1 m; they are also easily distinguished from other land flatworms by the characteristic hammer shape of their head. Bipaliines, which have their origin in warm parts of Asia, are invasive species, now widespread worldwide. However, the scientific literature is very scarce about the widespread repartition of these species, and their invasion in European countries has not been studied. Methods: In this paper, on the basis of a four year survey based on citizen science, which yielded observations from 1999 to 2017 and a total of 111 records, we provide information about the five species present in Metropolitan France and French overseas territories. We also investigated the molecular variability of cytochrome-oxidase 1 (COI) sequences of specimens. Results: Three species are reported from Metropolitan France: Bipalium kewense, Diversibipalium multilineatum, and an unnamed Diversibipalium 'black' species. We also report the presence of B. kewense from overseas territories, such as French Polynesia (Oceania), French Guiana (South America), the Caribbean French islands of Martinique, Guadeloupe, Saint Martin and Saint Barthelemy, and Montserrat (Central America), and La Reunion island (off South-East Africa). For B. vagum, observations include French Guiana, Guadeloupe, Martinique, Saint Barthelemy, Saint Martin, Montserrat, La Reunion, and Florida (USA). A probable new species, Diversibipalium sp. 'blue,' is reported from Mayotte Island (off South-East Africa). B. kewense, B. vagum and D. multilineatum each showed 0% variability in their COI sequences, whatever their origin, suggesting that the specimens are clonal, and that sexual reproduction is probably absent. COI barcoding was efficient in identifying species, with differences over 10% between species; this suggests that barcoding can be used in the future for identifying these invasive species. In Metropolitan south-west France, a small area located in the Department of Pyrenees-Atlantiques was found to be a hot-spot of bipaliine biodiversity and abundance for more than 20 years, probably because of the local mild weather. Discussion: The present findings strongly suggest that the species present in Metropolitan France and overseas territories should be considered invasive alien species. Our numerous records in the open in Metropolitan France raise questions: as scientists, we were amazed that these long and brightly coloured worms could escape the attention of scientists and authorities in a European developed country for such a long time; improved awareness about land planarians is certainly necessary

Obama chez moi! The invasion of metropolitan France by the land planarian Obama nungara (Platyhelminthes, Geoplanidae)

Background: Obama nungara is a species of land flatworm originating from South America; the species was recently described and distinguished from a similar species, Obama marmorata. Obama nungara has invaded several countries of Europe, but the extent of the invasion has not been thoroughly mapped. Methods: In this article, based on a five and a half-year survey undertaken by citizen science, which yielded 530 records from 2013 to 2018, we analysed information about the invasion of Metropolitan France by O. nungara. We also investigated the variability of newly obtained cytochrome c oxidase 1 (COI) sequences of specimens from France, Italy and Switzerland. Results: Obama nungara was recorded from 72 of the 96 Departments of Metropolitan France. The species is especially abundant along the Atlantic coast, from the Spanish border to Brittany, and along the Mediterranean coast, from the Spanish border to the Italian border. More than half of the records were from an altitude below 50 m, and no record was from above 500 m; mountainous regions such as the Alps, Pyrenees and Massif Central are not invaded. Local abundance can be impressive, with 100 of specimens found in a small garden. An analysis of our new COI sequences, combined with published sequences of specimens from several countries, confirmed that three clades comprise the species. The first clade, 'Brazil', is currently confined to this country in South America; the second clade, 'Argentina 2', was found in Argentina and in Europe, only in Spain; and the third, 'Argentina 1', was found in Argentina and in Europe, in Spain, Portugal, France, UK, Italy, Belgium, and Switzerland. This suggests that two clades of O. nungara from Argentina have invaded Europe, with one widely spread. Discussion: The present findings strongly suggest that O. nungara is a highly invasive species and that the population which has invaded several countries in Europe comes from Argentina. The wide dispersion of the species and its reported local abundance, combined with the predatory character of the species, make O. nungara a potential threat to the biodiversity and ecology of the native soil fauna in Europe, and probably the most threatening species of all invasive land planarians present in Europe

Hammerhead flatworms (Platyhelminthes, Geoplanidae, Bipaliinae): mitochondrial genomes and description of two new species from France, Italy, and Mayotte

Background: New records of alien land planarians are regularly reported worldwide, and some correspond to undescribed species of unknown geographic origin. The description of new species of land planarians (Geoplanidae) should classically be based on both external morphology and histology of anatomical structures, especially the copulatory organs, ideally with the addition of molecular data. Methods: Here, we describe the morphology and reproductive anatomy of a species previously reported as Diversibipalium “black”, and the morphology of a species previously reported as Diversibipalium “blue”. Based on next generation sequencing, we obtained the complete mitogenome of five species of Bipaliinae, including these two species. Results: The new species Humbertium covidum n. sp. (syn: Diversibipalium “black” of Justine et al., 2018) is formally described on the basis of morphology, histology and mitogenome, and is assigned to Humbertium on the basis of its reproductive anatomy. The type-locality is Casier, Italy, and other localities are in the Department of Pyrénées-Atlantiques, France; some published or unpublished records suggest that this species might also be present in Russia, China, and Japan. The mitogenomic polymorphism of two geographically distinct specimens (Italy vs France) is described; the cox1 gene displayed 2.25% difference. The new species Diversibipalium mayottensis n. sp. (syn: Diversibipalium “blue” of Justine et al., 2018) is formally described on the basis of external morphology and complete mitogenome and is assigned to Diversibipalium on the basis of an absence of information on its reproductive anatomy. The type- and only known locality is the island of Mayotte in the Mozambique Channel off Africa. Phylogenies of bipaliine geoplanids were constructed on the basis of SSU, LSU, mitochondrial proteins and concatenated sequences of cox1, SSU and LSU. In all four phylogenies, D. mayottensis was the sister-group to all the other bipaliines. With the exception of D. multilineatum which could not be circularised, the complete mitogenomes of B. kewense, B. vagum, B. adventitium, H. covidum and D. mayottensis were colinear. The 16S gene in all bipaliine species was problematic because usual tools were unable to locate its exact position. Conclusion: Next generation sequencing, which can provide complete mitochondrial genomes as well as traditionally used genes such as SSU, LSU and cox1, is a powerful tool for delineating and describing species of Bipaliinae when the reproductive structure cannot be studied, which is sometimes the case of asexually reproducing invasive species. The unexpected position of the new species D. mayottensis as sister-group to all other Bipaliinae in all phylogenetic analyses suggests that the species could belong to a new genus, yet to be described

Negative impacts of invasive predators used as biological control agents against the pest snail Lissachatina fulica: the snail Euglandina ‘rosea’ and the flatworm Platydemus manokwari

Since 1955 snails of the Euglandina rosea species complex and Platydemus manokwari flatworms were widely introduced in attempted biological control of giant African snails (Lissachatina fulica) but have been implicated in the mass extinction of Pacific island snails. We review the histories of the 60 introductions and their impacts on L. fulica and native snails. Since 1993 there have been unofficial releases of Euglandina within island groups. Only three official P. manokwari releases took place, but new populations are being recorded at an increasing rate, probably because of accidental introduction. Claims that these predators controlled L. fulica cannot be substantiated; in some cases pest snail declines coincided with predator arrival but concomitant declines occurred elsewhere in the absence of the predator and the declines in some cases were only temporary. In the Hawaiian Islands, although there had been some earlier declines of native snails, the Euglandina impacts on native snails are clear with rapid decline of many endemic Hawaiian Achatinellinae following predator arrival. In the Society Islands, Partulidae tree snail populations remained stable until Euglandina introduction, when declines were extremely rapid with an exact correspondence between predator arrival and tree snail decline. Platydemus manokwari invasion coincides with native snail declines on some islands, notably the Ogasawara Islands of Japan, and its invasion of Florida has led to mass mortality of Liguus spp. tree snails. We conclude that Euglandina and P. manokwari are not effective biocontrol agents, but do have major negative effects on native snail faunas. These predatory snails and flatworms are generalist predators and as such are not suitable for biological control

New and revised terrestrial flatworm taxa (Platyhelminthes: Tricladida: Terricola) of Australia and the Subantarctic Islands of New Zealand

Within the context of ongoing taxonomic revisions of terrestrial flatworms from Australia and the Subantarctic Islands of New Zealand: (1) a new subfamily of the Terricola, the Anzoplaninae, is erected, and some subantarctic terricolan taxa are reassigned to the new subfamily; (2) diagnoses for the subfamily Caenoplaninae and genus Coleocephalus are emended; (3) the new genera Anzoplana and Fyfea are erected; and (4) the new species Anzoplana trilineata is described. A lectotype is designated for Fyfea carnleyi (Fyfe)

Studies on the systematics and biogeography of terrestrial flatworms (Platyhelminthes: Tricladida: Terricola) of the Australian region

This study contributes to knowledge of the systematics and biogeography of terrestrial flatworms (Platyhelminthes: Tricladida: Terricola) of the Australian region. Terrestrial flatworms are carnivorous soil animals some of which are important commercially as biological control agents for the Giant African snail, or as introduced pest species preying upon earthworms in Europe. The total Australian terrestrial flatworm fauna is estimated to be in excess of 300 species. At present only 137 species are named of which three-quarters have been described solely from external features. Taxonomy of the Terricola is based on a combination of external features and internal anatomical characters and remains in a state of flux. Recent molecular studies of the Tricladida have cast doubt on the monophyly of the Terricola. The austral fauna is poorly known anatomically, and taxonomy of the group further hampered by the existence of numerous species complexes which are poorly understood. The objectives of this study are to locate and examine type and supplementary specimens, review the functional anatomy of austral terrestrial flatworms, identify reliable taxonomic characters and states, revise certain taxa, and develop identification keys. Furthermore, efforts have been made to determine the distribution of terricolan taxa of the region, identify the biogeographical components of the austral terricolan fauna and to identify the occurrence of introduced, threatening and vagrant terrestrial flatworms. Cladistic and molecular analyses of the taxa have not been undertaken. Fixation and parasite-induced artifacts can cause mis-interpretation of anatomical characters. A suite of taxonomic characters and states is used to provide standardized diagnoses for genera. Included in the characters assessed for their application in taxonomy are colour and pattern of markings, pharyngeal musculature, configuration of the nephridial ducts, dorsoventral distribution of testes, penis type, resorptive organs, and musculoglandular organs (adenodactyls). Seven types of the hitherto enigmatic adenodactyls are identified, some of which function to secrete sclerotins which form the cocoon wall. The mechanism of cocoon formation in the Terricola involves the deposition of sclerotins, derived from antral secretions or adenodactyls, on the outer pre-cocoon membrane. This mechanism differs from the generally accepted Triclad model based upon the Paludicola, in which the cocoon is formed from within the pre-cocoon membrane by shell-globule components of the vitellocytes. Representatives of all three terricolan families are present in the Australian region. A new subfamily is erected in the Geoplanidae, and of the twenty-seven genera considered in the study, ten are new. The problematic Australoplana complex is resolved into five genera and two sub-genera. Field keys are provided for the Terricola genera of the Australian region. This study supports the view that the Terricola are polyphyletic. Anatomical data suggest a possible phylogenetic relationship between some austral caenoplaninid and maricolan taxa. The Australian flatworms mostly occur within hyper-humid to sub-humid moisture regions, and are assigned to two principal families. In the Rhynchodemidae: Rhynchodeminae 45 species are accommodated within Cotyloplana, Digonopyla, Dolichoplana, Platydemus, Rhynchodemus and in new genera not considered in this study. In the Geoplanidae: Caenoplaninae, 80 species are assigned to Artioposthia, Australoplana, Caenoplana, Fletchamia, Lenkunya, Parakontikia, Reomkago, Tasmanoplana and additional new genera considered here. Four species are assigned to two new genera in a new subfamily of the Geoplanidae. Eight introduced species are identified, none of which at present poses an ecological or commercial threat in the Australian region. Two major flatworm faunal units are recognized in Australia. A northern element dominated by rhynchodemid genera including taxa which also occur in Papua-New Guinea - Irian Jaya and Indonesia. The southern element is dominated by geoplanid genera including some with currently poorly-defined Gondwanan affinities. These flatworm faunal units broadly accord with the Torresian (northern) and with the Tasmanian, South-western, and Bassian or Kosciuskan (southern) zoogeographic subregions

The Australian terrestrial flatworm fauna (Tricladida : Terricola)

Australian terrestrial flatworms number more than 137 known species of which 70% are described. The total fauna is estimated to be in excess of 300 species. The flatworms mostly occur within hyper-humid to sub-arid moisture regions, and are assigned to two principal families. In the Rhynchodemidae: Rhynchodeminae 35 species are accommodated in Cotyloplana, Digonopyla, Dolichoplana, Platydemus, Rhynchodemus, and new genera. The 84 species in the Geoplanidae: Caenoplaninae are assigned to Artioposthia, Caenoplana, Australoplana, Parakontikia, Reomkago, Fletchamia, Lenkunya, Tasmanoplana, and new genera. There are eight introduced species. Two major flatworm faunal units are recognised in Australia. A northern element dominated by rhynchodemid genera including taxa which also occur in Papua New Guinea - Irian Jaya and Indonesia. The southern element is dominated by geoplanid genera including some with as yet poorly defined Gondwanan affinities. These flatworm faunal units broadly accord with the Torresian (northern) and with the Tasmanian, South-western and Kosciuskan (southern) zoogeographic subregions