Methods of classifying nemerteans: an assessment

Phenetic, cladistic and phyletic methods of classifying animals are discussed with particular reference to nemerteans. It is concluded that phenetic (numerical) taxonomy is particularly inapplicable to any group of invertebrates for which well defined character differences are relatively few, whilst both the phenetic and cladistic methods fail through their fundamental assumption that convergent evolution is a rare occurrence. Terrestrial and freshwater nemerteans especially demonstrate convergent evolution in many ways; cladistic classifications proposed for these animals are therefore untenable. Convergence is shown to be a common occurrence in other nemerteans also. It is concluded that because the traditional phyletic approach does not implicitly assume that resemblances between organisms are more likely to be due to common ancestry than to convergence, it is far more likely to reveal true evolutionary relationships between taxa.     

Terrestrial nemerteans thirty years on

The present paper records changes in the reported distribution and abundance of terrestrial nemerteans during the last 30 years. New records are reported from the Isles of Scilly, Bermuda and New Zealand of species previously only known elsewhere. Human activity, especially the increased importation and exportation of plants between countries, has probably led to terrestrial nemerteans, as well as other organisms, becoming introduced into many new localities. However, the conservation outlook is not hopeful because of a global loss of appropriate habitat, particularly forests. Today three species (Geonemertes rodericana Gulliver, 1879, Pantinonemertes agricola Willemoes-Suhm, 1874 and Antiponemertes allisonae Moore, 1973) are feared to be extinct, and none of the extant species seem to be as abundant as they once were. Behavioural changes in Antiponemertes pantini (Southgate, 1954) under arid conditions may be associated with heavy gregarine infestations.     

THE EVOLUTION AND COMPARATIVE PHYSIOLOGY OF TERRESTRIAL AND FRESHWATER NEMERTEANS

(1) Parallel evolution from separate marine ancestors has given rise to two groups of genera of terrestrial nemerteans, representative of at least two different family stocks. They have colonized land by the direct route, moving up the shore on widely separated oceanic islands. The relationships between the known terrestrial and semi-terrestrial nemerteans, comprising seven genera and 15 species, are defined and their geographic distribution is discussed.
(2) In fresh water, three heteronemertean and nine hoplonemertean species are known. The hoplonemerteans arise from two different families representing two different evolutionary routes to fresh water, one by way of estuaries and the other via land. The distribution of freshwater nemerteans and the characters common to all species are discussed.
(3) Very little is known about the comparative physiology of land and freshwater nemerteans. However, information about the physiology of marine forms is combined with morphological differences from them found in terrestrial and freshwater species to suggest profitable lines of future research.
(4) Water relations pose a major problem for both terrestrial and freshwater nemerteans. The role of the cerebral organs, the blood and the excretory system in the osmotic physiology of some littoral nemerteans is beginning to be understood. These systems in land and freshwater forms have clearly distinctive features which need to be thoroughly investigated.
(5) Suggestions are given for valuable lines of investigation of the mucous glands, digestive physiology and general behaviour of terrestrial and freshwater nemerteans.     

Exploring the molecular diversity of terrestrial nemerteans (Hoplonemertea, Monostilifera, Acteonemertidae) in a continental landmass

We report the finding of terrestrial nemerteans of the family Acteonemertidae from the northern part of Spain. The specimens were studied using molecular data from the nuclear ribosomal 18S rRNA and the mitochondrial cytochrome c oxidase subunit I genes. The 18S rRNA data strongly suggest the presence of three species of terrestrial nemerteans in the Iberian Peninsula, an old landmass where terrestrial nemerteans had not previously been reported. These specimens originate from primarily undisturbed forests across a distance of c . 1000 km. Cytochrome c oxidase subunit I data also indicate the presence of multiple lineages of Iberian terrestrial nemerteans. Nonetheless, the pattern obtained from this marker is obscured most probably by deep genetic divergences. This molecular diversity, at least in some of the clades, suggests that the Iberian species are not the result of recent introductions, as proposed for other terrestrial nemerteans found in Europe. Our data also touch upon the question of a single origin of terrestriality in nemerteans, a hypothesis rejected by both data sets. Nevertheless, terrestriality seems to have had a single origin in the family Acteonemertidae.     

Food, Feeding Behavior and Feeding Ecology of Nemerteans

The feeding biology of nemerteans is reviewed, new informationis presented, and the role of nemerteans in natural communitiesis discussed and evaluated. Most nemerteans are carnivorous,the majority feeding on live, often specific, prey, while someare scavengers. Macrophagous feeding is found in the Palaeonemertea,Heteronemertea and some of the Hoplonemertea; the more specializedsuctorial feeding is limited to five of the hoplonemertean families,and suspension feeding (omnivorous diet) occurs only in thehighly specialized symbiotic monogeneric Bdellonemertea. Feeding adaptations seem to be related to the nature and employmentof the proboscis, the dilatability of the mouth, and the structureof the anterior part of the digestive tract. The transport ofdissolved organic materials from seawater has been demonstrated,but its role in nemertean nutrition is just being contemplated.Present meager information on predation has shown nemerteansto have actually and potentially large effects on prey populationsand thus on the communities in which they live. As the feedingbiology of only a relative handful of species has been studied,much basic comparative research is needed before we can corroborateor refute our present ideas concerning the ecological rolesof nemerteans.     

The Necessity of DNA Taxonomy to Reveal Cryptic Diversity and Spatial Distribution of Meiofauna,with a Focus on Nemertea

Meiofauna represent one of the most abundant and diverse communities in marine benthic ecosystems. However, an accurate assessment of diversity at the level of species has been and remains challenging for these microscopic organisms. Therefore, for many taxa, especially the soft body forms such as nemerteans, which often lack clear diagnostic morphological traits, DNA taxonomy is an effective means to assess species diversity. Morphological taxonomy of Nemertea is well documented as complicated by scarcity of unambiguous character states and compromised by diagnoses of a majority of species (and higher clades) being inadequate or based on ambiguous characters and character states. Therefore, recent studies have advocated for the primacy of molecular tools to solve the taxonomy of this group. DNA taxonomy uncovers possible hidden cryptic species, provides a coherent means to systematize taxa in definite clades, and also reveals possible biogeographic patterns. Here, we analyze diversity of nemertean species by considering the barcode region of the mitochondrial gene Cytochrome Oxidase subunit I (COI) and different species delineation approaches in order to infer evolutionarily significant units. In the aim to uncover actual diversity of meiofaunal nemerteans across different sites in Central America, COI sequences were obtained for specimens assigned here to the genera Cephalothrix, Ototyphlonemertes, and Tetrastemma-like worms, each commonly encountered in our sampling. Additional genetic, taxonomic, and geographic data of other specimens belonging to these genera were added from GenBank. Results are consistent across different DNA taxonomy approaches, and revealed (i) the presence of several hidden cryptic species and (ii) numerous potential misidentifications due to traditional taxonomy. (iii) We additionally test a possible biogeographic pattern of taxonomic units revealed by this study, and, except for a few cases, the putative species seem not to be widely distributed, in contrast to what traditional taxonomy would suggest for the recognized morphotypes.     

Aspects of the biology of Pantinonemertes californiensis,a high intertidal nemertean

I studied the distribution, feeding biology, and reproductive biology of Pantinonemertes californiensis, described as a semi-terrestrial nemertean, along the central California coast. At the sites used in this study, maximal tidal height is about 2 m, and P. californiensis typically occurred under boulders between 1.3 and 1.7 m tidal height. Worms fed primarily on the semi-terrestrial amphipod Traskorchestia traskiana. Distribution of nemerteans was similar to that of the prey, although prey extended higher on the beach than did the worms. Nemerteans were largest and most abundant at the site with highest abundance of T. traskiana and smallest and least abundant at the lowest prey abundance site. In laboratory feeding trials, nemerteans from the site with lowest prey abundance fed most readily. Non-reproductive nemerteans lived for at least a week when submerged in sea water; some prey died within a week of being submerged. Nemerteans only lived minutes when submerged in fresh water; 50% of prey lived 4.5 h. Eggs are approximately 90–100 μm in diameter and hundreds to thousands are shed per female. Larvae are planktonic and apparently planktotrophic, and are morphologically similar to other marine hoplonemertean larvae. At the sites studied life history characteristics of P. californiensis provided little evidence of adaptations to terrestrial life in these worms and were not helpful in elucidating the role of semi-terrestrial nemerteans in the evolution of terrestrial nemerteans.     

Ecological implications of the reproductive biology of symbiotic nemerteans

A major problem for parasites is to ensure that offspring will reach a correct host. Responses to this problem include several reproductive adaptations. This review paper shows that Carcinonemertes has several such adaptations, including increased number of gonads and offspring, synchrony of hatching with host larval hatching, facultative parthenogenesis, and mating. Other nemerteans show different, but comparably adaptive, reproductive modifications for parasitism. In contrast, reproductive biology of the commensal Malacobdella resembles that of free-living nemerteans, being timed to coincide with plankton blooms that are food for settling young.It has been thought that nemerteans found in association with other animals usually have only causal relationships with those animals, and that actual parasitism is rare among nemerteans. However, it is being found that parasitism is both well developed and more common among nemerteans than was previously thought. Nemerteas show in their reproductive biology that they are adapted to the selective factors that are important in their ways of life, and we should be able to predict the life styles of these worms by knowing their reproductive ecology.