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.     

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.     

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.     

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.     

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.     

The peculiar nemertean larva pilidium recurvatum belongs to Riserius sp., a basal heteronemertean that eats Carcinonemertes errans,a hoplonemertean parasite of Dungeness crab

A typical nemertean pilidium larva resembles a hat with ear flaps. But one type, called pilidium recurvatum, looks more like a sock, swimming heel first. This distinctive larva was discovered in 1883 off the coast of Rhode Island and subsequently found in plankton samples from other parts of the world. Despite the long time since discovery, and its significance in discussions of larval evolution, this larva remained unidentified even to the family level. We collected pilidium recurvatum larvae from plankton samples in Coos Bay, OR, and identified them as belonging to the heteronemertean genus Riserius based on juvenile morphology and DNA sequence data. Phylogenetic analysis suggests that two distinct types of pilidium recurvatum from Oregon represent two new species within this currently monotypic genus. We describe the morphology of pilidium recurvatum using confocal microscopy and compare it to that of the typical pilidium, discussing possible implications for larval feeding. We also report our surprising discovery that juveniles of Riserius sp. from Oregon prey on another nemertean, Carcinonemertes errans, an egg predator of Cancer magister (Dungeness crab), a commercially important species. We speculate that the species‐level diversity and geographic distribution of Riserius may be much greater than currently appreciated.     

Predation on soft-shell clams (Mya arenaria) by the nemertean Cerebratulus lacteus in Atlantic Canada: implications for control measures

The nemertean, Cerebratulus lacteus Verrill (Nemertinea: Heteronemertini), has been identified as an important threat to soft-shell clam (Mya arenariaL.) populations in Atlantic Canada. The biology of this species, however, is still largely unknown. Field and laboratory studies were undertaken in 1998 and 1999 in Prince Edward Island, Canada, to test certain control measures to reduce predation on soft-shell clam populations and to better describe the relationship between C. lacteus and M. arenaria. Field abundance of C. lacteus, M. arenaria and Nereis virens Sars were evaluated in relation to particular habitat modifications that were used as control measures. Sediment manipulations tested were: (1) addition of shells and (2) use of a hydraulic rake. No difference was observed on the abundance of C. lacteus, M. arenaria and N. virens after treatments were applied. In the laboratory, C. lacteus was shown to be an efficient predator of M. arenaria. Clam mortalities reached 100% in the presence of C. lacteus while 0% mortality was observed in its absence. A complementary set of experiments was carried out to see if the sympatric polychaetes N. virens and Glycera dibranchiata Ehlers had any impact on the relationship between C. lacteus and M. arenaria. N. virens showed no impact on C. lacteus predation on clams. The presence of G. dibranchiata significantly reduced the nemertean predation rate. Analysis of clam size selection revealed no significant preference by C. lacteus. Other experimental studies revealed that high predator densities did not impede predation on clams and that C. lacteus preferred soft-shell clams among other commercial bivalve species presented (Mercenaria mercenariaL., Crassostrea virginica Gmelin and Mytilus edulisL.). This study should provide a better understanding of the relationship between C. lacteus and M. arenaria and lead to the development of improved nemertean control measures.     

Nemertean larvae of the ototyphlonemertidae family in the plankton of Peter the Great Bay, Sea of Japan

Larvae of two nemertean species,Ototyphlonemertes (O.) martynovi Chernyshev, 1993 andOtotyphlonemertes (Norenburgia) sp., found in the neritic plankton of Peter the Great Bay, the Sea of Japan are described. The larvae of both species had two statocysts and a pair of eyes, but differed in the number of statoliths and in their tactile cirri. Changes of some traits during ontogenesis are discussed.     

A reassessment of the systematics and a proposal for the phylogeny of some cosmopolitan Lineus species (Nemertea)

Morphological, behavioural and biochemical characters of 9 cosmopolitan, benthic species of the nemertean genus Lineus are used both to clarify their taxonomy and to discuss their phylogenetic relationships. Data analysis shows that all fissiparous species of Lineus collected in the world seas up to date can be placed in a single species, L. sanguineus, with three recognisable subspecies, L. s. sanguineus, L. s. nigricans and L. s. pseudolacteus. These taxa were originally described as separate species mainly according to their geographic origin under the names L. sanguineus (shores of European seas), L. socialis (Atlantic North American shores), L. vegetus (Pacific North American shores), L. pseudolacteus (shores of the English Channel) and L. nigricans (Mediterranean coasts of Italy and France). Examination of the character value matrix suggests a phylogenetic tree developing from two ancestral branches. The first branch includes the two related species L. ruber and L. viridis, the second branch the three species L. longissimus, L. lacteus and L. sanguineus. This approach to the taxonomic and phylogenetic relationships of Lineus nemertean species is in agreement with the intra- and interspecies histocompatibility data in nemerteans: (i) grafts transplanted from donors to recipients of the same species succeed and grafts transplanted from donors to recipients of different species fail; (ii) grafts are rejected more rapidly when the donor and recipient species diverged earlier in evolution and, a contrario, grafts are accepted better when the donor and recipient species have been more recently isolated.     

Embryonic and larval development of a cold adapted Antarctic ascidian

Ova of the Antarctic ascidian Cnemidocarpa verrucosa were mature at 240–245 μm. At 0 to −1.5°C, embryos hatched as swimming tadpoles at 8 days from fertilization, which is close to the ages at which some Antarctic echinoderm and nemertean embryos hatch as blastulae. Comparisons of Antarctic and temperate ascidian larvae suggest that the ascidian’s development rate is affected by low environmental temperatures to about the same extent as embryos and larvae of an echinoid, nemertean, and calanoid copepods. The ascidian’s tadpoles were bright orange and large, >2 mm in length including tunic and >1.5 mm in length without tunic. The large and brightly colored tadpoles were conspicuous when swimming, which supports the hypothesis that larvae of C. verrucosa are chemically defended against predators. Metamorphosed juveniles were found in cultures within 16 days from fertilization, when some unsettled tadpoles still moved but were less active. The potential pelagic period may therefore be 16 or more days with 8 days as an unhatched embryo and up to 8 or more days as a tadpole. The resting metabolic rate of tadpole larvae was 15 pmol O₂ h⁻¹ individual⁻¹ which is equivalent to larval respiration rates in Antarctic echinoderms. A low resting metabolic rate suggests a potential mechanism for the extended larval lifespan in C. verrucosa.     

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 .     

Community Structure of Macrobenthic Invertebrates in Sandy Beaches of the Jordan Gulf of Aqaba,Red Sea

Six sandy beach stations along the Jordan Gulf of Aqaba were selected for macrofaunal examination during November 1980. Fifteen core samples (0.01 m² to a depth of 10–15 cm) were taken at each station; five at the high tide line, five at the low tide line, and five midway. Twenty-two taxa were encountered from all stations. Polychaetes were the most diverse group with seven taxa, followed by molluscs and crustaceans with six and five taxa respectively. Animal diversity, using the Shannon-Wiener Index, was generally low. Based on the biological index value of McCLOSKEY (1970), seven taxa were considered the community dominants. These are: the polychaetes, Glycera tesselata, Pisione remota, Saccocirrus sp., and Perinereis nuntia; the isopod Eurydice arabica; a nemertean and an oligochaete.     

The structure of the proboscis musculature in <Emphasis Type="Italic">Baseodiscus delineatus</Emphasis> (Delle Chiaje, 1825) (Heteronemertea) and the comparative analysis of the proboscis musculature in heteronemerteans

The proboscis musculature was studied in the nemertean Baseodiscus delineatus using confocal laser scanning and electron transmission microscopy. Three muscle layers were differentiated in the proboscis wall: the outer-longitudinal, the diagonal, and the inner-circular layer. The endothelium consists of two cell types: apical supportive cells with rudimentary cilia and subapical myocytes making up the inner-circular musculature of the proboscis. The supportive cells have thin processes attached to the basal extracellular matrix and their perikarya are spread over the apical surfaces of myocytes. The endothelium of B. delineatus is characterized by a folded basal layer of the extracellular matrix and by different heights of myocyte processes, giving an impression that the inner-circular musculature is multilayered. Comparative analysis shows that the diagonal musculature of Baseodiscus is not homologous to that of other heteronemerteans. An assumption is made that the inner-circular muscles have endothelial origin in all heteronemerteans.     

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.     

Food and feeding behavior of the hoplonemertean Oerstedia dorsalis

The monostiliferous nemertean Oerstedia dorsalis was collected from eelgrass (Zostera marina) beds located along the coast of New Jersey, and feeding responses to amphipods and isopods were observed in the laboratory. Tests with 46 worms showed that they fed suctorially on Ampelisca vadorum, Ampithoe longimana, Corophium acherusicum and C. tuberculata. Corophiids were preferred. Upon contact with an amphipod, the proboscis is everted and strikes the prey on the ventral side, immobilizing it in a few minutes. The worm probes the sternal region with its head and everts its proboscis one or more times during the process. The exoskeleton is eventually penetrated by the head, and the stomach is everted into the hemocoel as a flattened funnel-like structure. Peristaltic undulations of the body signify the suctorial action that removes the living contents from the exoskeleton. The actual feeding process (from head penetration to removal of the head) takes about 7 min. O. dorsalis is only the third species within the Prosorhochmidae for which the feeding behavior has been documented. The other two are terrestrial species, and are also suctorial.     

Effects of salinity, temperature and pH on the survival of the nemertean Procephalothrix simulus Iwata, 1952

The salinity, temperature and pH tolerance of Procephalothrix simulus Iwata, 1952, were experimentally studied. In hypo-media, the nemerteans could survive 96 h in 3.3‰ solution at 10 °C (median lethal salinity [LS₅₀] was not determined at this temperature), and 96 h LS₅₀ were 7.3‰ and 13.5‰ at 20 °C and 30 °C, respectively. In hyper-media, 96 h LS₅₀ values were 53.9‰, 47.1‰ and 41.4‰ at 10 °C, 20 °C and 30 °C, respectively. The trend of body weight changes in diluted media indicated that this nemertean is a volume regulator. During a 96-h exposure in media at 0 °C, worms were thanatoid but could recover if the temperature was gradually elevated to 20 °C. In thermal tolerance experiments, the nemertean survived 96 h in seawater of 30 °C, and worms suffered high mortalities when the temperature exceeded 32 °C. Present results suggest that the interaction of temperature and salinity on the lethal effects on P. simulus is significant (P < 0.05). Elevated temperature (range 10-30 °C) decreased the worm's solute tolerance, and elevated salinity (range 18-38‰) decreased the worm's thermal tolerance. The survival pH level for this nemertean ranged from 5.00 to 9.20.