Predation on snails by three species of triclad and its bearing on the distribution of Planaria torva in Britain

Dugesia polychroa, D. lugubris and Planaria torva preyed upon snails of differing morphologies and while the Dugesia spp. fed approximately in proportion to snail abundance, PL torva was more selective on Potamopyrgus jenkinsi. The incidence of feeding on four snail taxa by the Dugesia spp. was significantly correlated and differed strikingly from that of PI. torva where feeding was related to its annual life cycle. Large peaks of feeding coincided in the Dugesia spp. for the four snail types; this was attributed to snail vulnerability being increased by environmental disturbance. Comparison of feeding by Dugesia spp. in the presence and absence of PI. torva indicated that severe competition was occurring for food with PI. torva having a food refuge in P. jenkinsi. The distribution of PI. torva in Britain is largely confined to productive habitats in the Glasgow-Edinburgh and East Anglia regions. It is concluded that historical rather than ecological events have played the major part in its distribution to date. The strong association of PL torva with port areas and the canal system suggests that it is a recent introduction via the timber trade with Fennoscania.     

Diet of the leeches Erpobdella octoculata (L) and Helobdella stagnalis (L) in a lotic habitat subject to organic pollution

SUMMARY 1. A study of the diet of E. octoculata (L) and H. stagnalis (L) using a serological technique was carried out in a lot ic habitat subject to organic pollution and seasonal sewage fungus growth.
2. The results indicate that E. octoculata consumed predominantly Chironomidae and Oligochaeta, while H. stagttalis had a less pronounced prey preference, consuming rather similar proportions of chironomids, oligochaetes, Asellus and gastropod snails.
3. At enriched sites, E. ocioculata consumed relatively more chironomids and fewer oligochaetes, while H. stagnalis took more gastropods and fewer chironomids.
4. Seasonal differences were noted in the diet of both leeches with chironomids predominating chiefly from April to August, and molluscs, oligochaetes and Asellus at other times of the year. Intra-specific differences (on a weight-age basis) were only seen for E. octoculata , small forms consuming chiefly oligochaetes and larger ones concentrating on chironomids.     

The diet ofEnoplus brevis (Nematoda) in a supralittoral salt marsh of the North Sea

The gut content of nearly 2000 specimens ofEnoplus brevis was quantitatively analysed.E. brevis is an opportunistic omnivore. Main food categories found were cyanobacteria, diatoms, oligochaetes, nematodes, and rotifers. With a few exceptions, the diet of females and males was identical. Predominantly, juveniles fed on cyanobacteria and adults on animal prey. Experiments revealed that living oligochaetes are attacked and preyed upon. The quantities of the food categories eaten byEnoplus varied according to their ambient abundance, both temporarily and spatially on a small scale. At least the nematode prey was strongly selected: some species were significantly preferred and others avoided.     

The food niches of the invasive Dugesia tigrina (Girard) and indigenous Polycelis tennis Ijima and P. nigra (Müller) (Turbellaria; Tricladida) in a Welsh lake

The invasion of Llyn Coron in North Wales by the American immigrant Dugesia tigrina resulted in the almost entire displacement of the native Polycelis tenuis and P. nigra populations. Because competition for food is the most important factor controlling and regulating populations of British lake-dwelling triclads, the diets of the invasive and native triclads in the lake were examined in an attempt to explain the successful invasion.A serological technique, the precipitin test, identified the gut contents of field-collected triclads. Niche breadth, electivity and niche overlap indices were used in analysis of the data. A broad food niche was recorded for all the triclad species. Oligochaetes predominated in the diet of D. tigrina followed by Asellus and chironomids, in almost equal proportions, snails and caddisflies, with mayflies scarcely eaten. Both Polycelis species fed heavily on oligochaetes followed by Asellus. P. tenuis ate slightly higher proportions of chironomids and caddisflies than snails and mayflies, whilst P. nigra consumed similar proportions of these four prey taxa. Gammarus, scarce in Llyn Coron, and cladocerans were not eaten by the Polycelis species and by only a few D. tigrina.To-date there has been no apparent shift in the diet of the Polycelis species as a consequence of the invasion by D. tigrina. In the absence of any evidence for the partitioning of food on the basis of prey size or condition, the considerable overlap in the diets of D. tigrina and the Polycelis species suggests the potential for severe inter-specific competition for food. The superior exploitation by D. tigrina of the available food resource requires explanation.     

The feeding biology of the immigrant triclad Phagocata woodworthi in Loch Ness, Scotland

Laboratory feeding experiments and a serological examination of the gut contents of field-collected triclads indicated that Phagocata woodworthi Hyman feeds heavily on oligochaetes. Its realized food niche also includes Asellus , caddisflies, molluscs, chironomids, stoneflies and mayflies. It is concluded that Ph. woodworthi is a considerable threat to the British triclad fauna, because of the considerable overlap in diet with native species, its fast reaction time to damaged prey, and its polypharyngeal state.     

The reaction time of leech and triclad species to crushed prey and the significance of this for their coexistence in British lakes

1. Three species of leeches, Erpobdella octoculata, Glossiphonia complanata and Helobdella stagnalis, and four species of triclads, Polycelis nigra, P. tennis, Dugesia polychroa and Dendrocoelum lacteum, commonly coexist on stony shores in productive British lakes. All species are food limited and there is much overlap in their diet. For both leech and triclad communities, coexistence of species is through the occurrence of food refuges. Leeches are more successful than triclads at capturing live prey, whereas both groups feed on damaged prey, comprising incapacitated, live or dead animals that are leaking body fluids. If triclads are better than leeches at exploiting damaged prey, this could be a mechanism for their coexistence. 2. Laboratory experiments investigated the comparative speeds at which leeches and triclads responded to crushed prey. Young and adult predators were offered a crushed specimen of the oligochaete Tubifex tubifex, the snail Lymnaea peregra, the crustacean Asellus aquaticus or the chironomid Chironomus sp., and their reaction times recorded. These four prey groups constitute the main diet of the predators in the field. Only D. polychroa and D. lacteum showed a significantly different reaction time between young and adults to crushed prey, and the reason for this is unclear. All predators, except H. stagnalis and D. polychroa, showed a difference in reaction time to the four types of prey, presumably a consequence of differences in both the ‘quality’ and ‘concentration’ of the different prey fluids, and there were some differences between predators in their speed of reaction to the same prey type. The following sequence, from fastest to slowest, in general reaction time to prey was obtained: E. octoculata, D. polychroa, P. tenuis, D. lacteum, P. nigra, H. stagnalis and G. complanata. 3. The location of the damaged food by the predators can be explained partly in terms of their foraging behaviour, with E. octoculata, D, polychroa and P. tenuis exhibiting a more seek-out strategy than other species which have a more sit-and-wait behaviour, and partly on the level of sophistication of their chemosensory system used to detect leaked prey fluids. This system is highly developed in triclad species but poorly developed in leeches. 4. In a second type of experiment in which prey, L. peregra, A. aquaticus or Chironomus sp., were offered at different time intervals after crushing to H, stagnalis and P. tenuis, few predators fed on food crushed for 24 h or longer, although a few leeches fed on Chironomus crushed for up to 72 h. 5. It is concluded that coexistence of leech and triclad species on stony shores in lakes is assisted by partitioning of food on a damaged or live basis.     

Feeding on gastropods by lake-dwelling Polycelis in the absence and presence of Dugesia polychroa (Turbellaria, Tricladida)

SUMMARY. Using a serological technique, the extent of feeding upon snails by Polycelis tenuis was compared in habitats lacking and containing Dugesia polychroa which is the major triclad predator of snails. It was found that P. tenuis ate significantly more snails when D. polychroa was absent. It was concluded that in the field interspecific competition for snails between these two triclad species was a more powerful process than any social feeding by P. tenuis on snails captured by D. polychroa .
Feeding on snails as a group by P. tenuis and P. nigra in the habitat where the two triclads co-existed was of a similar magnitude and this was also true of their feeding on the common snail species and genera. Approximately 10% of the triclads gave a positive reaction to snail antisera.
Feeding on snails by P. nigra in the three habitats lacking D. polychroa was similar in extent and did not differ from that of P. tenuis . It was concluded that P. nigra would be similarly affected by the presence and absence of D. polychroa as shown for P. tenuis .
No definitive seasonal pattern was detected in the incidence of feeding upon snails by either Polycelis species but this may have been obscured by seasonal changes in the detection period of snail antigens due to temperature effects. There was some evidence that such feeding increased during the period when small snails were present in the population.     

A comparative study of the food niches of lake-dwelling triclads and leeches

The food niches of four species of triclads and three species of non-parasitic leeches living in the littoral zone of British lakes of different trophic status were investigated, over one year, using a serological technique. Antisera against ten potential prey groups were employed. The basic data were adjusted to compensate for differences in predator size and seasonal changes in field temperatures. Data on two of the leech species have been presented elsewhere, but comparisons of the diet of all seven predator species are made in this paper. Much overlap in diet between the various predator species occurs, but, with one exception, each of the genera has a major food resource. The Polycelis spp. feed extensively on oligochaetes, Dugesia polychroa on molluscs, Dendrocoelum lacteum on Asellus, Erpobdella octoculata on chironomids and Glossiphonia complanata on molluscs. Helobdella stagnalis is a generalist feeder. Problems of coexistence of the leech species, and of the leech and triclad species are discussed.     

A comparative assessment of temporal variation in diet of powan, Coregonus lavaretus (L.), from Loch Lomond and Loch Eck, Scotland, U.K.

Stomach contents of powan caught throughout the year from Loch Lomond and Loch Eck were examined. Adult powan in Loch Lomond fed mainly on planktonic Cladocera, switching to benthic food during December-April. Bosmina coregoni was the principal planktonic food with Daphnia hyalina an important late summer and autumn prey. Loch Eck powan fed throughout the year on chironomid larvae and Pisidium spp. from the benthos and ingested large amounts of non-food material. Mean dry weights of food in stomachs were correlated with water temperature in Loch Lomond but not in Loch Eck and were higher in powan from Loch Eck over most of the year.     

Food utilization and niche overlap in three species of lake-dwelling leeches (Hirudinea)

The diets of Erpobdella octoculata, Glosiphonia complanata and Helobdella stagnalis on the stony littoral zone of eutrophic Crose Mere were examined over two years using the precipitin test. The abundances of food organisms were measured. For all leech species, juvenile mortality was greatest in locations where a low ratio of prey abundance to recruited leech abundance occurred, and the intensity of feeding by Erpobdella and Helobdella fell slightly in late summer when large numbers of juveniles of these species were present; these suggest the involvement of food in juvenile mortality, Seasonal changes in feeding intensity were similar for all leech species, with the exceptions outlined above and for Glossiphonia in April when feeding by brooding adults ceased, at which time the severity of any interspecific competition would have been reduced. Within a leech species, there was little change in the food spectrum as the leeches grew. Niche breadth was lowest for Erpobdella and highest for Helobdella. Niche overlap was greatest between Glossiphonia and Helobdella, and there was a trend for overlap values to be lower between Glossiphonia and Erpobdellu than between Helohdellu and Erpohdellu. Erpobdella fed mainly on chironomids and oligochaetes, Glossiphonia on molluscs, oligochaetes and chironomids, and Helobdella on oligochaetes, chironomids, Asellus, molluses and amphipods, all in decreasing order of importance. It is suggested that coexistence in the leech species may be due in part to Erpobdella and Glossiphonia having 'food refuges' in chironomids and molluscs, respectively, with Helobdellu being more of a general predator.     

Food preferences of tanypodinae larvae (Diptera: Chironomidae)

Analyses of the gut contents of larvae of three species of Tanypodinae, collected from four sites in north-east England and Scotland, invariably showed small particles (1–10 µm²), predominantly of detritus, to be more abundant than large particles (10–100 µm²), mostly algae and diatoms. No animal remains, such as carapaces, head capsules and chaetae, or vascular plant fragments were found in any of the intestines. In contrast, food-choice and growth experiments suggested that animal food (particularly Tubificidae and Chironominae larvae) was important in the tanypod diet. In the food-choice experiments, six different foods offered to small (4–7 mm long) and to large (8–12 mm long) larvae of Procladius choreus Meigen, were chosen in the following order: live Tubificidae spp.; detritus; live Chironominae larvae; a mixture of algae and diatoms; live Ostracoda spp.; and live Cladocera spp. However, when the results of the individual choices were examined, it was clear that the small larvae had consumed greater amounts of the small types of food (i.e. detritus, algae and diatoms) in comparison to the large larvae. In the growth experiment, those larvae of P. choreus reared solely on Tubificidae, detritus or Chironominae larvae thrived, while those fed on algae and diatoms, ostracods or cladocerans grew more slowly and fewer survived the experimental period.It was concluded that although Tanypodinae larvae are primarily predators with definite food preferences in ideal foraging conditions, in adverse conditions they will utilize a range of available stand-by foods, the most important of which appears to be detritus.     

The realized food niches of three species of stream-dwelling triclads (Turbellaria)

SUMMARY. 1. The diet of Phagocata vitta, Crenobia alpina and Polycelis felina in a Welsh stream was examined, using a serologicat technique, at monthly intervals over 18 months.
2. For each triclad species, dietary composition changed little with time or increasing body size, and overall feeding intensity was greatest in summer and least in winter.
3. Alt species fed most heavily on oligochaetes, chironomids and Gammarus. Phagocata vitta fed lightly on Plecoptera, Trichoptera and Ephemeroptera whilst the other two triclad species fed substantially on the first two of these prey categories.
4. Food niche breadth indices indicated a wide diet for all species with some specialization in some months. Electivity indices showed positive electivity for oligochaetes and chironomids by all species and for Plecoptera by C. alpina and P. felina .
5. Diets overlapped and none had a food refuge: therefore the potential for interspecific competition for food exists.     

Food partitioning by lake-dwelling triclads and glossiphoniid leeches: field and laboratory experiments

The triclads Polycelis tenuis and Dugesia polychroa and the glossiphoniid leeches Glossiphonia complanata and Helobdella stagnalis are abundant on the stony shores of productive British lakes. All species are food limited and there is considerable overlap in the diets of these triclads and leeches. This paper investigates interactions between the two groups using field and laboratory experiments to try to identify the mechanism of their co-existence. Triclad and leech numbers were manipulated inside experimental enclosures, mathced by controls, erected on the stony shore of an eutrophic English lake. Increasing the numbers of P. tenuis and D. polychroa prior to the reproductive season in spring resulted in a significant decrease in the numbers and body size of G. complanata and H. stagnalis compared with control populations in the summer months, and vice versa. However, increases and decreases were temporary with a readjustment of numbers and body size to control levels in the autumn after reproduction had ceased. It is suggested that increasing the numbers of either group elevated the severity of both intra- and interspecific competition for food. The condition of prey may, in part, determine the strength of competition, and this was examined in laboratory experiments in which different densities and ratios of P. tenuis and H. stagnalis were offered either live of recently crushed Asellus aquaticus. In monospecific controls, growth rates of P. tenuis were greater when fed on crushed than live Asellus, but there was no significant difference in the growth of H. stagnalis fed either live or crushed prey. In mixed cultures of predators, P. tenuis and H. stagnalis were the superior competitors when fed on crushed and live Asellus, respectively. However, when competitive pressure was low, at low densities of predators, the presence of H. stagnalis in mixed cultures fed on live prey was beneficial to the growth of P. tenuis. These results are explained in terms of the greater ability of triclads to detect damaged prey, leaking body fluids, due to their sophisticated chemosensory system, and the ability of leeches to capture live prey due to the presence of suckers. It is concluded that co-existence of the two groups in British lakes is assisted by the partitioning of food on a live or damaged basis.     

First results of a study of the feeding habits of <Emphasis Type="Italic">Synodontis</Emphasis><Emphasis Type="Italic">comoensis</Emphasis> (Siluriformes: Mochokidae) in a West African river (Comoé River,Comoé National Park,Côte d’Ivoire)

The feeding habits of Synodontis comoensis (Siluriformes, Mochokidae) have been studied for the first time. Specimens caught in the Comoé River (Comoé National Park, Côte d’Ivoire) had fed on different types of food including insects (Diptera, Ephemeroptera, Hymenoptera, and Isoptera), molluscs (gastropods and bivalves), oligochaetes, and macrophytes. The food items most frequently found in the stomach contents were chironomid larvae, oligochaetes, vegetable detritus, fruit, and gastropods. Correlation studies based on the index of preponderance of food items revealed noticeable variation in S. comoensis diet composition with fish size and season. Preferred food items of young fish were oligochaetes in the rainy season and vegetable detritus in the dry season. Food items preferred by larger fish were fruit and gastropods in the rainy season and chironomid larvae in the dry season.     

Continuing dispersal of freshwater triclads (Platyhelminthes; Turbellaria) in Britain with particular reference to lakes

1. A survey of lakes in Anglesey and Snowdonia, Wales, in the summer of 1997 recorded changes in the triclad fauna which had occurred since earlier studies in the 1950s and in 1973. Two native species, Polycelis tenuis and Dugesia polychroa, and two immigrants, Planaria torva and especially Dugesia tigrina, have increased their range. The triclad fauna has remained unchanged with time in eleven out of sixteen lakes in Snowdonia, in contrast to only one out of fourteen lakes on Anglesey. This supports the hypothesis produced in the 1950s, that the triclad fauna of the ion-poor, unproductive lakes in Snowdonia would change little with time, being maintained by ecological interactions, whereas the ion-rich, productive lakes on Anglesey, from which native Dugesia spp. and Dendrocoelum lacteum are historically absent, would increase in triclad diversity. 2. An examination of national records for the distribution of D. tigrina and Pl. torva indicated their continuing dispersal in both still and running waters in mainland Britain, with the former species having the greater frequency of occurrence. 3. The reasons for this may be that D. tigrina is (a) more easily dispersed, (b) a more opportunistic, catholic feeder, (c) a more vigorous competitor, (d) able to reproduce asexually and (e) able to prey to some extent on other triclad species, as shown in the laboratory but not yet confirmed in the field. The absence of D. tigrina and Pl. torva from unproductive lakes may be explained in terms of low temperature and feeding mode, and a low standing crop of gastropods, respectively. 4. Seven surveys (1961–1997) of the triclad fauna (six species) of Colemere, England, revealed that D. tigrina was confined to one small area of the littoral zone until the mid-1980s, after which it spread fairly rapidly to occupy the entire shore by 1997. Its expansion has been to the detriment of all the native species, particularly P. tenuis, except Dd. lacteum. The possible extinction of Pl. torva may be linked to a numerical reduction in snails, particularly Potamopyrgus jenkinsi. 5. Dugesia tigrina has an adverse effect on the native triclad fauna in lakes, particularly when only a few species are present. This has been explained in terms of the availability of a wider variety of food. The presence of a larger number of triclad species denies the immigrant food items, particularly crustaceans and snails, the food refuges of Dd. lacteum and Dugesia spp., respectively. However, the long-term study of Colemere has shown that more diverse triclad faunas may also be vulnerable to this immigrant. The effect of Pl. torva on native triclads remains uncertain.     

Gut evacuation of walleye pollock larvae in response to feeding conditions

Gut residence times of first-feeding walleye pollock larvae were measured at 6°C in continuous and discontinuous feeding regimes. Larvae fed tagged copepod prey evacuated their guts more quickly in continuous feeding as compared to the discontinuous feeding treatment. The mean gut residence time was estimated to be 5.0 h for larvae feeding continuously. Gut evacuation by larvae fed tagged prey and then isolated without food (discontinuous treatment) was slower and more variable, with an estimated gut residence time exceeding 8.0 h. Field and laboratory observations suggest that the larval fish gut may be modeled as an intermittent plug-flow reactor (PFR) in response to diel feeding patterns. The cyclical nature of gut dynamics has implications for gut content analyses and the estimation of daily food rations.     

Diet and niche overlap of southern populations of brill Scophthalmus rhombus and turbot Scophthalmus maximus

The diets of adult brill Scophthalmus rhombus and turbot Scophthalmus maximus from the Portuguese coast relied mostly on fishes. There was a higher diversity of food items compared to their northern counterparts, and several of the identified prey are the first records of these species, including a brown alga, echinoderms, nematodes, oligochaetes, gastropods, bivalves and various fish species. The diet of the two species was significantly different and niche overlap was low.     

Damage released prey alarm substances or predator odours? Risk assessment by an aquatic oligochaete

Although the abilities of prey to detect and respond to chemical substances associated with a predator have been widely reported, the factors promoting the evolution of responses to prey alarm cues vs. predator odours are still vague. In this article, we combined field research with laboratory experiments to explore which chemical substance associated with predator activity (predator odour, conspecific or heterospecific alarm substances) induces defence responses in the aquatic oligochaete Stylaria lacustris, which is vulnerable to common littoral predators. The field results indicated that predators injure the oligochaetes and a great proportion, up to 45% of individuals in the population, were found to be damaged. The results of the laboratory experiments revealed that chemical odours from damselfly larvae feeding on S. lacustris did not induce the defence response in the oligochaetes. On the contrary, oligochaetes detected and responded to alarm substances from damaged conspecifics alone and substances from damaged cladoceran Daphnia magna. We discussed conditions favouring the responses to damage released prey alarm cues instead of predator odours in Stylaria lacustris. Our data suggest that the selection of responses to alarm cues from damaged prey vs. predator odours may be dependent on three factors: (1) non-species-specific predation, (2) divergence of food niche of the different stages of the predator and (3) complex food web with multiple predators. Handling editor: S. Declerk