A new genus and species of monostiliferoidean nemertean (Nemertea: Enopla) found on an egg mass of the anomuran decapod Paralithodes camtschatica

A new genus and species of monostiliferoidean enoplan nemertean from Alaska is described and illustrated. The nemertean, Alaxinus oclairi gen. et sp. nov. , was found on the egg mass of a red king crab, Paralithodes camtschatica.     

A new species of Tetrastemma (Nemertea: Enopla: Monostiliferoidea) from Ria de Foz, north-western Spain, found living in the mantle cavity of the bivalve mollusc Scrobicularia plana

A new species of monostiliferoidean nemertean, Tetrastemmafozensis , from the mantle cavity of the bivalve mollusc Scrobicularia plana , in the Ría de Foz, north-western Spain, is described and illustrated. Other nemerteans from the mantle cavity of Scrobicularia collected in Poole Harbour, southern England, are provisionally identified as conspecific with the Ria de Foz material, but show certain anatomical differences whose taxonomic significance cannot at present be assessed.     

Hubrecht,Macfarlane, Jensen and Willmer: on the nature and testability of four versions of the nemertean theory of vertebrate origins

Four authors (A. A. W. Hubrecht, J. M. Macfarlane, D. D. Jensen and E. N. Willmer) have argued that nemerteans are the proximal invertebrate ancestors of vertebrates. While all these authors accept certain homologies (i.e. vertebrate pituitary and notochord with nemertean proboscis invagination and proboscis sheath or mesoderm), there is considerable variation among them in the derivation of the vertebrate nervous system, sense organs and foregut apertures. For example, Hubrecht suggests that the vertebrate nervous system developed from a solid dorsal nerve found in some nemerteans; Macfarlane suggests it developed by fusion of the left and right solid brains of nemerteans; Jensen suggests that the vertebrate tubular nervous system developed by the backward extension of the nemertean frontal organ and fusion with the nemertean dorsal and ventral brain ganglia. Modern EM, histochemical and biochemical methods can test empirically the four theories. Research testing these theories will also test the more popular acraniate theory, since that theory implies that vertebrate head structures evolved de novo in chordates and that no homologies should exist between vertebrate and nemertean head structures.     

Mutation and Selection Cause Codon Usage and Bias in Mitochondrial Genomes of Ribbon Worms (Nemertea)

The phenomenon of codon usage bias is known to exist in many genomes and it is mainly determined by mutation and selection. To understand the patterns of codon usage in nemertean mitochondrial genomes, we use bioinformatic approaches to analyze the protein-coding sequences of eight nemertean species. Neutrality analysis did not find a significant correlation between GC12 and GC3. ENc-plot showed a few genes on or close to the expected curve, but the majority of points with low-ENc values are below it. ENc-plot suggested that mutational bias plays a major role in shaping codon usage. The Parity Rule 2 plot (PR2) analysis showed that GC and AT were not used proportionally and we propose that codons containing A or U at third position are used preferentially in nemertean species, regardless of whether corresponding tRNAs are encoded in the mitochondrial DNA. Context-dependent analysis indicated that the nucleotide at the second codon position slightly affects synonymous codon choices. These results suggested that mutational and selection forces are probably acting to codon usage bias in nemertean mitochondrial genomes.     

A new genus and species of freshwater monostiliferous hoplonemertean (Nemertea, Enopla) from the People's Republic of China

A new genus and species of freshwater monostiliferous hoplonemertean, Limnemertes poyangensis gen. et sp. nov., from Poyang Lake, People's Republic of China, is described and illustrated. The taxon is compared and contrasted with previously described freshwater hoplonemerteans. This is the fourth species of freshwater nemertean to be described from China and the first recorded from Poyang Lake.     

Laboratory observations on the feeding behavior and feeding rate of the nemertean Procephalothrix simulus

The feeding behavior of the nemertean Procephalothrix simulus and the effects of extrinsic factors on the feeding rates of this nemertean were experimentally studied. Procephalothrix simulus showed a typical macrophagous feeding strategy and consumed various food items. When capturing freshwater Tubifex sp., the nemerteans successfully ingested prey in all attacks, and they did not evert the proboscis in 25% of capture events. When capturing marine Saccocirrus gabrillae, the nemerteans failed to consume prey in more than half of the attacks, and they always everted the proboscis. There was a positive relationship between nemertean body weight and the rate of successful attacks, and a negative relationship between nemertean body weight and the duration of feeding events. The feeding rate of P. simulus increased when the temperature was raised from 5 degrees C to 30 degrees C but was significantly inhibited at 32 degrees C. Food intake was significantly reduced in media diluted to a practical salinity of 20 and 10 and in medium with the salinity elevated to 45. Dark conditions induced higher food intake, but prey density had no significant effect on feeding rate. These results suggest that P. simulus is a predator successfully adapted to the variable environmental conditions of the intertidal habitat.     

Preliminary investigation of Vibrio alginolyticus-like bacteria associated with marine nemerteans

A marine bacterium provisionally identified as Vibrio alginoliticus, which is associated withtetrodotoxin production, was isolated from severalspecies of marine nemerteans. The presence of thisbacterium strongly supports the potential use oftetrodotoxins by nemerteans for protection againstingestion by predators or as an agent in their ownpredatory activities. Tetrodotoxins may facilitatethe ability of the nemertean proboscis to subdue preyduring capture. One commensal nemertean speciesappears not to possess the bacterium. This may bedue, in part, to its non-predatory life style.     

Phylogenetic Relationships among Higher Nemertean (Nemertea) Taxa Inferred from 18S rDNA Sequences

We estimated the phylogenetic relationships of 15 nemertean (phylum Nemertea) species from the four subclasses Hoplo-, Hetero-, Palaeo-, and Bdellonemertea with 18S rDNA sequence data. Three outgroup taxa were used for rooting: Annelida, Platyhelminthes, and Mollusca. Parsimony and maximum-likelihood analyses supported the monophyletic status of the Heteronemertea and a taxon consisting of hoplonemerteans and Bdellonemertea, while indicating that Palaeonemertea is paraphyletic. The monophyletic status of the two nemertean classes Anopla and Enopla is not supported by the data. The unambiguous clades are well supported, as assessed by a randomization test (bootstrapping) and branch support values.     

Status of the Nemertea as predators in marine ecosystems

The ecology of nemertean predators in marine ecosystems is reviewed. Nemerteans occur in most marine environments although usually in low abundances. Some species, particularly in intertidal habitats, may reach locally high densities. During specific time periods appropriate for hunting, nemerteans roam about in search of prey. Upon receiving a stimulus (usually chemical cues), many nemertean species actively pursue their prey and follow them into their dwellings or in their tracks. Other species (many hoplonemerteans) adopt a sit-and-wait strategy, awaiting prey items in strategic locations. Nemerteans possess potent neurotoxins, killing even highly mobile prey species within a few seconds and within the activity range of its attacker. Most nemertean species prey on live marine invertebrates, but some also gather on recently dead organisms to feed on them. Heteronemerteans preferentially feed on polychaetes, while most hoplonemerteans prey on small crustaceans. The species examined to date show strong preferences for selected prey species, but will attack a variety of alternative prey organisms when deprived of their favourite species. Ontogenetic changes in prey selection appear to occur, but no further information about, e.g. size selection, is available. Feeding rates as revealed from short-term laboratory experiments range on the order of 1–5 prey items d^–1. These values apparently are overestimates, since long-term experiments report substantially lower values (0.05–0.3 prey items d^–1). Nemerteans have been reported to exert a strong impact on the population size of their prey organisms through their predation activity. Considering low predation rates, these effects may primarily be a result of indirect and additive interactions. We propose future investigations on these interactive effects in combination with other predators. Another main avenue of nemertean ecological research appears to be the examination of their role in highly structured habitats such as intertidal rocky shore and coral reef environments.     

The reproductive cycle of the entocommensal nemertean Malacobdella arrokeana and its symbiosis with the geoduck Panopea abbreviata

Abstract. The reproductive cycle of the entocommensal nemertean Malacobdella arrokeana inhabiting Panopea abbreviata from San José Gulf, Patagonia, Argentina, and the nemertean–host relationship were investigated. The mature nemertean population showed a sex ratio of 1:0.64, with females predominating. The number of individuals of M. arrokeana per host varied from 1–60 individuals (mean±SD=3.7±9.4 mm). Different stages of gonadic development were observed in individual nemerteans. Spawning peaks occurred during late summer and early spring. Sexually mature individuals of both sexes were present year round, possibly indicative of a continuous breeding period. Our results suggest that M. arrokeana has no cannibalistic habits and confirm its entocommensal symbiosis with P. abbreviata .     

DNA barcoding reveals a cryptic nemertean invasion in Atlantic and Mediterranean waters

For several groups, like nemerteans, morphology-based identification is a hard discipline, but DNA barcoding may help non-experts in the identification process. In this study, DNA barcoding is used to reveal the cryptic invasion of Pacific Cephalothrix cf. simula into Atlantic and Mediterranean coasts. Although DNA barcoding is a promising method for the identification of Nemertea, only 6 % of the known number of nemertean species is currently associated with a correct DNA barcode. Therefore, additional morphological and molecular studies are necessary to advance the utility of DNA barcoding in the characterisation of possible nemertean alien invasions.     

A new species of commensal hoplonemertean from Australia

A new series of hoplonemertean from Australia is described and named Gononemertes australiensis. The morphology of this nemertean, which lives commensally in the ascidian Pyura pachydermatina (Herdman) variety intermedia Herdman, is described and illustrated and its taxonomic placing discussed.     

Temporal variation in abundance of the egg predator Carcinonemertes epialti (Nemertea) and its effect on egg mortality of its host,the shore crab,Hemigrapsus oregonensis

An outbreak of the nemertean, Carcinonemertes epialti, was observed on Hemigrapsus oregonensis during October, 1982 at Campbell Cove, Bodega Harbor, California. Mean worm intensity (296 worms/crab) was the highest recorded for this nemertean egg predator on H. oregonensis. During the outbreak, male crabs were found to harbor more worms than both non-ovigerous and ovigerous females. Crab egg mortality was substantial; 83% of the ovigerous females experienced 75–100% brood mortality. The seasonal peak in worm abundance coincided with the seasonal low in crab reproduction at this locality. A method for estimating the impact of C. epialti on H. oregonensis natality was developed using crab size and fecundity, and worm prevalence and intensity. For a non-outbreak sampling period, a mean of 5.6% egg mortality was experienced by infested crabs for the period selected. Thus, brood mortality during the outbreak was much greater than that experienced at non-outbreak periods. Heavy fishing pressure on some commercially important crab species has been suggested as a possible factor inducing worm outbreaks and facilitating their continued persistence. These observations suggest that fisheries are not necessarily responsible for the outbreaks of nemerteans on commercially important crab species. However, fishing pressure may still be a sufficient condition to promote nemertean outbreaks.     

Feeding rates of the nemertean Prosorhochmus americanus (Hoplonemertea) on two species of gammaridean amphipods

Abstract. The intertidal hoplonemertean Prosorhochmus americanus is a common inhabitant of the fouling community of rock jetties of the southeast coast of the United States. We undertook a laboratory investigation of the feeding rate of this nemertean, which is a suctorial predator of amphipod crustaceans that co‐occur in abundance in the fouling community. While submerged in water (simulating high tide), worms fed on the tube‐building amphipods Jassa falcata and Corophium cf. insidiosum at rates of 0.19 amphipods nemertean^?1 d^?1 (n=10) and 0.26 amphipods nemertean^?1 d^?1 (n=14), respectively. These predation rates were not significantly different (two‐tailed t‐test, p>0.05), and are similar to those estimated in laboratory studies of other suctorial nemerteans. Many nemerteans are typically more active at night, and indeed, adults of P. americanus consumed more individuals of J. falcata during dark periods than during light periods (χ² analysis, p<0.05). However, no difference in consumption of individuals of C. cf. insidiosum was observed in dark versus light. We attribute these contrasting results to differences in tube‐building behavior exhibited by these two species of amphipod under laboratory conditions. Our results and those of other laboratory investigations suggest that nemerteans that prey on amphipods feed at a rate of ～0.2 prey items nemertean^?1 d^?1, but under natural conditions this rate may not be obtained because of limited feeding time, longer foraging distances, and emigration of prey from regions of high nemertean activity.     

FMRFamide-like immunoreactivity and arylamidase activity in turbellarians and nemerteans--evidence for a novel neurovascular coordinating system in nemerteans

The tetrapeptide Phe-Met-Arg-Phe-NH2 (FMRFamide) has been immunolocalized in the nervous systems of seven species of Turbellaria and four species of Nemertea. The 11 species represent all the major turbellarian and nemertean taxa, and illustrate most of the various life styles found in these animals. The FMRFamide-like reactivity coincides with histochemically demonstrable arylamidase activity in the nervous systems. It is suggested that the FMRFamide-like reactivity demonstrates the presence in these lower invertebrates of one or more biologically active peptides, analogous to those of higher invertebrates and chordates and acting as putative neurotransmitters and coordinators of growth, maturation and muscular activities. The arylamidases occurring with the peptides are probably an integral part of these peptide-mediated control systems. The nemertean vascular system is especially rich in arylamidases and is believed to be concerned primarily with peptidergic control of bodily functions, rather than with transport of metabolites.     

Ecological studies of the nemertean fauna in an estuarine system of the northwestern Gulf of Mexico

The distribution and abundance of nemerteans in the brackish-water lakes of Sea Rim State Park, Texas, near the Louisiana border, were studied and compared with other macrobenthos during one year. Six of 93 macrobenthic species collected were nemerteans (0.9% of the total number of specimens). Only one species of nemertean, Carinoma sp., was consistently present. This species is the most ubiquitous and, probably, the most abundant nemertean in the estuarine systems of the Texas coast. Carinoma sp. was collected at Sea Rim from a salinity range of 0–21 ppt and at other Texas estuaries from 2–26 ppt. Preliminary experiments with Carinoma sp. as predator and as prey indicated that it feeds on polychaete worms and in turn is fed upon by white (Penaeus setiferus) and brown (P. aztecus) shrimp.     

Biology of a hoplonemertean from the branchial chambers of the pinnotherid crab Zaops (= Pinnotheres) ostreum

The pea crab, Zaops ostreum, a symbiont in the mantle cavity of the American oyster (Crassostreavirginica), is itself a host to a symbiotic nemertean. Thisnemertean belongs to the genus Carcinonemertes, and issimilar to or possibly the same as C. pinnotheridophila,known from another sympatric pinnotherid crab, Pinnixachaetopterana. Living in mucous sheaths within the gillchambers of female crabs, the life cycle of the nemertean isintimately synchronized with and dependent on the crab's reproduction. A total of 138 pea crabs were collected andexamined for nemerteans during the summers of 1965 to 1967 atBeaufort, North Carolina, U.S.A.; of these 97 (70%) were mature(5th stage) female crabs. Mature females harbored thenemerteans, and had a prevalence of 39.2%. A maximum of 20worms was found in the gill chambers of a single crab, 9 ofwhich were the red, eyeless, mature female worms, along with 11colorless males. Worm egg sacs were attached to the pleopods ofovigorous crabs; a maximum of 293 sacs was recorded in anindividual crab with 7 mature females. At about 110 eggs persac, this translates to a potential production of >32 200nemertean larvae. Since female Z. ostreum may live for 2to 3 yr and produce two broods of eggs per year (beyond thefirst year), the reproductive potential of the symbionts may beeven greater. Gills of crabs with more than one mature femalewere usually covered by the worm sheaths, and were frequentlydamaged (gnarled, torn, silted, etc.). The sternum of infestedcrabs was often damaged due to the projection of worm sheathsthrough it to the subabdominal region. There the egg sacs weredeposited on the pleopods to develop along with the crabembryos.