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.     

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.     

Behavioral responses to fish kairomones and autotomy in a damselfly

The threat-sensitivity hypothesis predicts that prey species assess and adjust their behavior in accordance with the magnitude of the threat posed by a predator. A largely overlooked characteristic of a prey that will affect its sensitivity to predators is its history of autotomy. We studied threat-sensitive behavior to fish kairomones in larvae of Ischnura elegans damselflies, which had undergone autotomy, from a fishpond and from a fishless pond. In agreement with their higher perceived risk, larvae from the fishpond showed fewer rigid abdomen bends, foraged less and walked more slowly than larvae from the fishless pond. In line with their higher vulnerability to predators, larvae without lamellae spent less time foraging than larvae with lamellae. There was a decrease in swimming activity in the presence of fish kairomones except for larvae with lamellae from the fishless pond. This may reflect differences in vulnerability of larvae without lamellae between pond types. Such context-dependent responses in activity to kairomones should be kept in mind when evaluating the ability of a prey to recognize kairomones.     

Does body size difference in the leeches Glossiphonia complanata (L.) and Helobdella stagnalis (L.) contribute to co-existence?

The effect of predator and prey body size on the feeding success of the British lake-dwelling leeches Glossiphonia complanata and Helobdella stagnalis was examined in the laboratory, and any involvement of size difference between the leeches in allowing coexistence in the field assessed. G. complanata breeds in advance of H. stagnalis and maintains a body size advantage throughout their annual life-cycle. In experiments, conducted at 14 °C and a photoperiod of 16 hrs L: 8 hrs D, three size classes of leeches of each species were each exposed to each of three size classes of each of five prey species, viz. Tubifex sp., Chironomus sp., Asellus aquaticus, Lymnaea peregra and Potamopyrgus jenkinsi. For each prey species, three different types of experiments were performed: one leech exposed to four prey individuals; four leeches of the same species with sixteen prey; and two leeches of each species with sixteen prey. In the first experiment, all sizes of G. complanata were capable of feeding on all sizes of the prey types offered; the same was true for H. stagnalis with exceptions of feeding on large A. aquaticus and large L. peregra. For both species, but especially for G. complanata, there was a trend within each size class of leech for decreasing proportions of fed leeches with increasing prey size, and within each size class of prey for an increasing proportion of fed leeches with increasing leech size; however there were several exceptions to these trends. Both leeches fed extensively on Tubifex sp. but there were significant differences in the proportions feeding on other prey types; G. complanata fed more on A. aquaticus and the two snail species, and less on Chironomus, than H. stagnalis. The effect of increasing the number of leech individuals from one to four individuals, of the same or mixed species, had little effect on the proportion of leeches which had fed. It is concluded that large G. complanata will have access to large individuals of certain prey taxa denied H. stagnalis, which may lessen the intensity of interspecific competition.     

The incidence of predation on lake-dwelling leeches

SUMMARY. 1. Erpobdella octoculata (L.), Glossiphonia complanata (L.) and Helobdella stagnate (L.) were eaten readily by larval and adult dytiscid beetles, larval caddisflies and Sialis , nymphal odonatans and various fish in laboratory experiments.
2. A serological technique showed that very few lake-collected specimens of these predatory taxa, plus triclads, leeches and stone fly nymphs, had been feeding on leeches.
3. A literature survey of lake studies on fish diet indicated that usually less than 6% of individuals with gut contents had eaten leeches. Few detailed records of waterfowl feeding on leeches have been documented.
4. It is concluded that predation pressure on lake-dwelling leeches is light, and is more severe from fish than invertebrates, less on populations inhabiting stony shores than in vegetation, and greater on the erpobdellid than the two glossiphoniid species.     

Behaviour, growth and survival of redfish larvae in relation to prey availability

Growth, survival and condition of redfish larvae Sebastes spp., reared in the laboratory (0, 500 1500 and 4500 prey l^-1) were highest in the 1500 prey l^-1 treatment. Significantly lower larval growth and survival in the 4500 prey l^-1 treatment corresponded with lower prey bite: orient ratios in later weeks, suggesting that larvae were unable to forage efficiently at high prey densities. While these prey densities are higher than those reported in the field, naturally co-occurring Atlantic cod Gadus morhua larvae require higher prey densities when reared under similar conditions in the laboratory. These data suggest that prey availability may not be as limiting to redfish as for other commercially important marine species.     

The occurrence of metacercariae of the trematode, Cyathocotyle opaca (Wisniewski), in three species of lake-dwelling leeches

SUMMARY. 1. The leeches, Erpobdella octoculata, Glossiphonia complanata and Helobdella stagnalis , were collected at monthly intervals over a 2-year period from a eutrophic English lake, and examined for the presence of metacercariae of the trematode. Cyathocotyle opaca.
2. Only one specimen of each of H. stagnalis and G. complanata was recorded with the parasite. Seasonal changes in prevalence levels in E. octoculata were similar for each of the three cohorts identified during the study period. Values, low in young leeches recruited in the autumn, rose slowly until the following spring, and then increased more rapidly till the autumn, when the cohort died out. Mean intensity and relative density showed similar patterns of seasonal change to that of prevalence.
There was no evidence of parasite induced mortality in E. octoculata. Attainment of sexual maturity did not differ between infected and uninfected leeches.     

The influence of temperature on the life cycle and occurrence of three species of lake-dwelling leeches (Annelida: Hirudinea)

The influence of constant temperatures on the growth, maturation and reproduction of field-collected and F, generation Helobdella stagnalis, Glossiphonia complanata and Erpobdella octoculata was investigated. Leech material from a lowland, productive lake and from an upland, unproductive lake was cultured at each of the temperatures: l·5,5·5,8, 10·5, 13,16, 19 and 23°C. Field-collected and F₁ generation H. stagnalis and E. octoculata matured and produced eggs and young over the range 10·5–23°C, whereas the range for egg production was 8–23°C for field-collected G. complanata, F, generation leeches failing to mature at any temperature due to an additional limiting factor being involved.
Optimal temperatures for cocoon/egg production were observed for E. octoculata, but egg production was similar at all temperatures for G. complanata and H. stagnalis. In contrast, the viability of eggs was greater at certain temperatures for the last two species but was similar at all temperatures for E. octoculata. Differences in egg production between field-collected and F, generation leeches are discussed. Rising temperatures accelerated rates of egg hatching, and of growth and maturation of leeches. The observed differences in survival of leeches at the various temperatures are discussed. Ecotypic adaptation to temperature within the small geographical area of Britain may not occur. It is concluded that though temperature may be a contributory factor determining distribution and abundance in British lakes, other causative factors, such as the occurrence and abundance of food organisms, are more likely to be implicated.     

Predation on leeches in a weedy pond

SUMMARY. 1. A range of sizes of Erpobdelta octoculata fL.), Glossiphonia complanata (L.) and Helobdella stagnalis (L.) were introduced to a small weeded pond harbouring invertebrate and amphibian predators. Control leeches were kept in protective cages. Subsequently, gut contents of potential predators were tested against three anti-leech sera, using the precipitin test. Five of fifteen species tested had fed on leeches; Agabus, Pyrrhosoma, Aeshna, Sympetrum and larval Trilurus.
2. The size distributions of introduced and surviving leeches were compared. Predation pressure had been greatest on smaller leeches and this may have accounted for the observed differential mortality between leech species.
3. It is concluded that predation may cause significant mortality in leech populations in weeded ponds, with severity determined by the types and abundance of predators present.     

Distribution and diel migration of macroinvertebrates within dense submerged vegetation

1. We studied vertical and horizontal distribution of macroinvertebrates within a dense stand of Chara spp. in Lake Krankesjön, southern Sweden. Invertebrates were sampled at three depths within the vegetation and at three distances from the vegetation edge during day and night in July and August. Corresponding samples of oxygen content of the water were taken.
2. The densities (number of invertebrates per unit plant biomass) of most invertebrate taxa were generally lower in the upper layers of the vegetation than in the layers close to the sediment. The densities of several taxa ( Asellus aquaticus , Cloëon sp. and Polycentropodidae), as well as total density of invertebrates, were higher at the edge than in the innermost parts of the macrophyte stand, whereas snail densities generally were highest at the innermost sites.
3. Densities of A. aquaticus , Chironomidae larvae and Helobdella stagnalis generally increased at night. These taxa appear to undertake a diel vertical migration within the vegetation, towards, or even down to the sediment in daytime and up into the vegetation, in some cases to the vegetation surface, at night. Factors underlying the diel vertical migration are discussed, as are their ecological consequences.     

A plant surface mutation mediates predator interference among ladybird larvae

Abstract.  1. We investigated mechanisms causing predator–predator interference between fourth instar Hippodamia convergens larvae foraging for pea aphids on pea plants, Pisum sativum , with a wild-type wax bloom, and the lack of such interference between larvae foraging on pea plants with a reduced-wax bloom caused by the single gene mutation wel .
2. Observations showed that behavioural interactions between larvae were not affected by wax phenotype. Specifically, larvae did not encounter one another more frequently on normal-wax peas as may have been predicted because reduced ability by coccinellids to attach to normal-wax plant surfaces could restrict them to foraging on only some parts of these plants.
3. In a controlled bioassay on normal-wax peas, H. convergens larvae avoided leaflets previously exposed to another larva. On reduced-wax peas, this effect was not detected.
4. In microcosm experiments, inter-predator interference in terms of prey consumption occurred on normal-wax peas, but not on reduced-wax peas. The interference on normal-wax peas occurred whether two H. convergens larvae were placed on a pea aphid-infested, normal-wax plant simultaneously or sequentially.
5. We conclude that the observed inter-predator interference is not as a result of direct physical contact, but rather arises because of (i) inhibition of foraging by chemical trails left by other larvae, (ii) the inability of larvae to access portions of the normal-wax plants creating aphid refugia, or (iii) a combination of these factors.     

Epibionts of Asellus aquaticus(L.) (Crustacea, Isopoda): an SEM study

1. This study presents a qualitative and quantitative survey of epibionts infesting two populations of the freshwater isopod Asellus aquaticus (L.). Using scanning electron microscopy, the prevalence, mean intensity, abundance, spatial preferences and distribution of organisms on various external surfaces were ascertained.
2. A diverse community, consisting primarily of protozoa and rotifers, utilised A. aquaticus as a substrate organism. Every individual A. aquaticus supported some form of epizoic life. Peritrich species comprised 89.2% of all epibionts. Carchesium polypinum, Pseudocarchesium aselli, Pseudocarchesium asellicola, Pseudocarchesium simulans and Opercularia hebes dominated the epifauna.
3. Highest epibiont burdens occurred on the mouthparts, the first few ventral segments and the gills and largely comprised the above peritrich species. Most epibionts exhibited some degree of site preference on the host. A distinct gill epifauna existed. Apart from Acineta tuberosa and Vorticella rotunda , which had equal prevalences on dorsal and ventral surfaces in one culture, most species largely avoided the dorsal surface.
4. Epibionts exhibited varying degrees of adaptation to life on a living substrate. They ranged from facultative species such as A. tuberosa , which are capable of colonising abiotic substrata, to Gymnodinoides aselli whose life cycle relies on interaction with A. aquaticus . Highly specialised species tended to exhibit the most pronounced and consistent site restriction.
5. Asellus aquaticus offers a highly tractable system for further study of substrate species–epibiont relationships.     

Waterborne metals impair inducible defences in Daphnia pulex: morphology, life-history traits and encounters with predators

1.  Inducible defences may be temporary and favoured where predation is intermittent and have been demonstrated in several invertebrates and vertebrates when prey detect chemical cues (kairomones) released by predators. Daphnia pulex (a water flea) exposed to Chaoborus (midge larvae) kairomones produce small neckteeth on the dorsal surface of the head as a defence against this gape-limited predator and survive better in the presence of Chaoborus . Recent studies have shown that waterborne copper (Cu) impairs the induction of neckteeth which could lead to lower survival.
2.  Here, we examined the effects of Cu on morphological changes and shifts in life-history traits in D. pulex exposed to kairomone from Chaoborus americanus . We exposed D. pulex mothers to chemical cues of C. americanus fed on either D. pulex neonates or on brine shrimp Artemia salina , the same Chaoborus cues combined with an environmentally relevant concentration of copper (10 μg L⁻¹), or dechlorinated tap water. We examined several morphological characteristics of neonates and life-historical characteristics of adults as well as assessing survival of neonates by staging encounters with predators.
3.  Neonates from mothers exposed to kairomone plus copper had fewer and shorter neckteeth than neonates from mothers exposed to kairomone alone. Moreover, neonates exposed to Cu had lower survival during encounters with predators than neonates exposed to kairomone without Cu.
4.  Adult female Daphnia exposed to kairomones released more neonates within the first 24 h of brood release and emptied their brood pouches quicker than mothers not exposed to kairomones, irrespective of the presence of Cu.
5.  Impairment by metals of morphological defences in zooplankton could lead to a decline in population density and alter community structure.     

用ELISA方法检测稻田捕食者对摇蚊的捕食作用

应用酶联免疫吸附（ELISA）双抗体夹心法,研究了稻田19种常见捕食性天敌对中性昆虫摇蚊的捕食作用。用摇蚊作抗原免疫雄性大白兔获得抗血清,用中和法、双向琼脂扩散实验及交叉反应对所制备的抗血清作特异性检测,抗体反应表明制备的抗血清对摇蚊抗原具有较高的特异性。测定了19种捕食者捕食摇蚊的临界吸收值。在检测的19种捕食者中,有13种捕食了摇蚊,占被检测捕食者种数的67.89％。ELISA阳性反应率最高的是在早稻前期采集的褶管巢蛛,阳性率高达50％,其次是晚稻中期采集的拟水狼蛛,其阳性率为40％。ELISA方法敏感,能快速检测捕食者对猎物的捕食作用及确定节肢类捕食者如蜘蛛对水稻害早控制作用大小,作为一种有效实验工具,可有助于发展水稻害虫综合管理理论。     

The abundance and trophic biology of Clarias albopunctatus Nichols & LaMonte, 1953 (Osteichthyes: Clariidae) in a tropical floodriver basin

The abundance, food and feeding biology of Clarias albopunctatus was studied over a period of 17 months in the lower River Anambra, Nigeria. The catfish was more abundant in late dry season (January–March) and early rainy season (April–June) than in other periods of the year in the small-sized Akwu pond than in other habitats. Feeding intensity was higher at night and at dawn (20.00–05.00 h) than during the day (08.00–17.00 h) in the 151–200 mm TL size class and during the rainy season (April–September). There was no sex-dependent variation in feeding intensity. Insects were the predominant food, followed by crustaceans. Of primary importance were Chironomidae (mainly Chironomus spp. and Tanypus sp.), Odonata nymphs, Dytiscus sp., mosquito larvae and pupae, Gyrinus sp., Daphnia sp., Ostracoda and Tilapia fry. Qualitative food composition and food richness were size, but not sex – or season-dependent. Diet breadth increased with catfish size and during the rainy season. The feeding behaviour of the catfish include foraging, shovelling and surface feeding. Abundant food and ability to switch to more available items enable C. albopunctatus to maintain its abundance in the River Anambra. This revised version was published online in July 2006 with corrections to the Cover Date.     

An energetic evaluation of a predator–prey (leech–amphipod) couple in Montezuma Well, Arizona, USA

1. The highly stable conditions in Montezuma Well, Arizona, have contributed to a very close predator–prey relationship between the endemic leech, Erpobdella montezuma , and its endemic amphipod prey, Hyalella montezuma , which forms nearly 90% of the leech diet.
2. Using a simple bioenergetic model, it was determined why a benthic opportunistic foraging strategy, typical of most other erpobdellids, is inferior to the specialized foraging behaviour utilized by E . montezuma and why reproduction by the leech is restricted during certain times of the year.
3. Our model demonstrated that to attain a biomass in excess of 2000 mg (maximum field size) during periods of low amphipod densities in the winter E . montezuma must feed on alternative prey types and have a foraging activity < 4 h per day.
4. Leech reproduction 4–6 months prior to the onset of winter food shortages of prey increases juvenile and adult survivorship with highest reproductive success for E . montezuma occurring in June–July.
5. Rather than being an obligate forager on H . montezuma, E . montezuma exploits a periodically abundant amphipod during most of the year in addition to alternate prey types during the winter thereby achieving an overall higher rate of energy gain than could be achievable by other closely related species of erpobdellids utilizing an opportunistic foraging approach.     

Temperature effects on chemical signalling in a predator–prey system

SUMMARY 1. We investigated the effect of temperature on chemical signalling in a predator–prey model system (planktivorous fish and Daphnia galeata ). Life-history changes in Daphnia in response to chemical cues (kairomones) derived from fish have become a paradigm for chemically induced anti-predator defences.
2. As temperature can affect both predator and prey, we carried out two experiments to disentangle these effects. In order to test for temperature effects on the predator, we kept prey at a single temperature and exposed them to kairomones from fish exposed to two different temperatures. Daphnia exhibited a higher intrinsic rate of population increase ( r ) when exposed to fish kairomones produced at high rather than low temperature. Assuming a positive correlation between r (because of an earlier maturation and/or increased clutch sizes) and kairomone concentration, our results suggest that kairomone production increases with rising temperature.
3. In the second experiment, to study the influence of temperature on the prey, Daphnia were kept at two different temperatures and exposed to fish kairomones produced at one constant temperature. We found no interaction between the effects of fish kairomone and temperature on Daphnia life history, suggesting that temperature does not directly alter life-history responses to fish kairomones.
4. Our results suggest that temperature influences Daphnia life history through its effects on fish kairomone concentration, but that temperature does not affect the strength of the response of Daphnia to the presence of fish.     

Prey‐driven behavioral habitat use in a low‐energy ambush predator

Food acquisition is an important modulator of animal behavior and habitat selection that can affect fitness. Optimal foraging theory predicts that predators should select habitat patches to maximize their foraging success and net energy gain, likely achieved by targeting areas with high prey availability. However, it is debated whether prey availability drives fine‐scale habitat selection for predators. We assessed whether an ambush predator, the timber rattlesnake (Crotalus horridus), exhibits optimal foraging site selection based on the spatial distribution and availability of prey. We used passive infrared camera trap detections of potential small mammal prey (Peromyscus spp., Tamias striatus, and Sciurus spp.) to generate variables of prey availability across the study area and used whether a snake was observed in a foraging location or not to model optimal foraging in timber rattlesnakes. Our models of small mammal spatial distributions broadly predicted that prey availability was greatest in mature deciduous forests, but T. striatus and Sciurus spp. exhibited greater spatial heterogeneity compared with Peromyscus spp. We found the spatial distribution of cumulative small mammal encounters (i.e., overall prey availability), rather than the distribution of any one species, to be highly predictive of snake foraging. Timber rattlesnakes appear to forage where the probability of encountering prey is greatest. Our study provides evidence for fine‐scale optimal foraging in a low‐energy, ambush predator and offers new insights into drivers of snake foraging and habitat selection.     

Prey behaviour and size-selective predation by fish

SUMMARY 1. We investigate the role of differential activity of chiro-nomids as an explanation for the size selection of small larvae by fish.
2. In the laboratory, pumpkinseed sunfish (Lepomis gibbosus L.) selectively consumed large larvae of Chironomus tentans Fab. when no tube-building materials were available. Small larvae were selectively consumed when a mud substrate was provided but there was no difference in predation rates on large and small larvae in sand.
3. Small larvae spent more time out of their tubes than large larvae in the presence of fish, which may explain the selection for small larvae in mud. Large larvae apparently compensated for decreased foraging activity in the presence of fish by increasing activity when fish were absent. Visibility of large larvae inside tubes may account for their increased mortality in sand.
4. Our results suggest that differential activity is important in explaining the size-selective mortality observed in the field.     

Sub-lethal effects of disturbance on a predatory net-spinning caddisfly

1. The sub-lethal effects of hydrologic disturbances on stream invertebrates are poorly understood, but integral to some models of how disturbances influence population and community dynamics. Carnivorous larvae of a net-spinning caddisfly, Plectrocnemia conspersa , have a strong predatory impact in some streams. Their silken nets, however, are vulnerable to high flow disturbance and the consequent destruction of nets could reduce predatory impacts and have life history consequences.
2. In a laboratory experiment, we manipulated the frequency of disturbances that destroyed the nets of P. conspersa , in the presence and absence of potential prey. Animals were housed individually and each trial lasted 8 days. We estimated net size, cumulative mass of silk produced, net allocation (net mass expressed as a proportion of body mass), per capita prey consumption and growth or mass loss of larvae.
3. In the absence of prey, increased disturbance frequency was accompanied by increased loss of body mass, a reduction of net size and an increase in the cumulative mass of silk produced. At the highest disturbance frequency, larvae eventually gave up producing nets. The ratio of net mass to body mass decreased with increasing disturbance, suggesting a trade-off in the allocation of resources, with a decreasing proportion of resources available for foraging. In the presence of prey, increased disturbance frequency was accompanied by a reduction in per capita prey consumption. Although foraging success offset the costs of silk production, growth rate decreased with increasing disturbance and could eventually lead to reduced body size and fecundity of adults.
4. These sub-lethal effects suggest that hydrologic disturbances could impose metabolic costs and reduce foraging efficiency of this predator. Thus, disturbances may reduce predator impact on prey populations and reduce predator population size without any direct mortality or loss of individuals.