The effects of feeding regime on the growth and reproduction of the medicinal leech Hirudo medicinalis

1. The feeding frequency, the size of meals, the number of meals required to attain reproductive maturity and the number of meals taken between iteroparous reproductive bouts were determined in the laboratory under optimal conditions for the medicinal leech Hirudo medicinalis fed exclusively on mammalian (bovine) blood. In addition the number of bouts of reproduction and the numbers of cocoons and hatchlings per cocoon produced were determined.
2. The average time for H. medicinalis to reach reproductive maturity at 20°C was 289 days, at an average wet biomass of 8143 mg with two–nine separate bouts of cocoon production. The number of meals to first reproduction was 8.9 (mean meal size of 3066.7 mg), with a significant correlation between total mass of blood ingested and the numbers of reproductive bouts and number of cocoons produced. Mean lifetime cocoon production per individual was 12.43, with 3.9 hatchlings per cocoon.
3. The significant positive relationships between ingestion, fecundity and developmental rate observed support the hypothesis that declining abundances of field populations of H. medicinalis are the result of lower available energy for growth, reflecting leeches now feeding predominantly on amphibian blood of lower energetic value than mammalian blood.     

Effects of chronic gamma irradiation on reproduction in the earthworm Eisenia fetida (Oligochaeta)

Eisenia fetida were exposed continuously to (60)Co gamma radiation during two generations (F(0) and F(1)). Adult F(0) reproduction capacity (i.e., number of cocoons produced, hatchability and number of F(1) hatchlings) in controls and at five dose rates (0.18, 1.7, 4, 11 and 43 mGy/h) was measured over a 13-week exposure period. Survival, growth and sexual maturation of F(1) hatchlings were observed for 11 weeks. F(1) adults were exposed for a further 13 weeks to determine their reproduction capacity. There was no radiation-induced effect on the cocoon production rate in either F(0) or F(1). For F(0), hatchability of cocoons produced during the first 4 weeks was reduced to 60% at 43 mGy/h (98% in controls), and none of the cocoons produced at 5-13 weeks hatched. At 11 mGy/h the cocoon hatchability was reduced to 25% at 9-13 weeks. In addition, the number of hatchlings per hatched cocoon was reduced at 11 and 43 mGy/h. Correspondingly, at these dose rates, the total number of F(1) hatchlings per adult F(0) was significantly lower than in the control. This number was also reduced at 4 mGy/h, but the effect was of borderline significance. For adult F(1), the hatchability of cocoons at 11 mGy/h was reduced to 45-69% during the 13-week exposure period. The number of hatchlings (F(2)) per cocoon and the total number of F(2) individuals produced was also reduced. However, and in contrast to the results observed for F(0), hatchability increased with time, suggesting a possible acclimatization or adaptation of the F(1) individuals. In conclusion, chronic irradiation reduced the reproduction capacity of E. fetida, but extensive exposure periods (13 weeks) were needed for these effects to be expressed. The lowest dose rates at which an effect was observed were 4 mGy/h in F(0) and 11 mGy/h in F(1).     

Scheme for dispersion analysis for evaluating heritability of reproductive traits in medicinal leech Hirudo medicinalis L

ANOVA designs for estimating heritabilities of fecundity traits in Hirudo medicinalis including batch size, the number of juveniles per cocoon, and juvenile weight. Accounting for reproduction mode of this species, different types of kinship were identified, which were taken into account in the ANOVA designs: juveniles from one batch and one cocoon were considered respectively full sibs and polyzygotic twins. Variation components were analyzed in the following kinship groups: for batch size, in full sibs; for the number of juveniles per cocoon, between sibships, full sins, and replicates; and for juvenile weight, in full sibs and polyzygotic twins. Using these designs, heritabilities of basic reproductive traits of H. medicinalis were obtained: for batch size, h2 = 0.35-0.40, H2 = 0.40-0.45; for the number of juveniles per cocoon, h2 = 0.33-0.36, H2 = 0.36-0.39; and for juvenile weight, h2 = 0.40-0.44, H2 = 0.44-0.48.     

The structure of a natural population of the medicinal leech, Hirudo medicinalis, at Dungeness, Kent

SUMMARY 1. A hot branding method was used to mark medicinal leeches ( Hirudo medicinalis L.) from five samples (four in 1984, one in 1985) at agravel pit at Dungeness in Kent. Analysis of capture-recapture data by the Fisher-Ford and Jolly methods gave similar results and indicated a population in excess of 10000 (0.112 leeches per m²) in 1984 and 1985. Successive estimates were each higher than the last suggesting a thriving community.
2. The population structure was skewed towards the lower weight classes. This may indicate good recruitment of hatchlings, as also indicated by the presence of cocoons, or the failure to capture heavier satiated or gravid leeches because of their relative unresponsiveness to the sampling method involving water disturbance, or their complete absence from the water during cocoon laying. Gravid leeches were always above 2 g in weight. Recaptured marked leeches revealed that they migrated throughout the site, indicating active foraging for hosts.     

The influence of body size on immediate reproductive success in Dugesia gonocephala (Tricladida,Paludicola)

The individual reproductive output of the stream-dwelling flatworm Dugesia gonocephala was investigated. Various measures of reproductive success were related to body size. (I) For the first 30 days in the laboratory small individuals produced no cocoons, individuals of intermediate size produced unfertilized cocoons and large individuals usually produced fertilized cocoons. (II) In individuals that produced a cocoon, no correlation was found between the number of cocoons produced in one month and body size. (III) Large individuals, however, produced larger cocoons. This was not due to the fact that unfertilized cocoons were smaller. (IV) Large cocoons tended to contain more young. (V) The average size of young hatching from large cocoons was larger. (VI) Large individuals produced their first cocoon soon after their arrival in the laboratory and seemed to have a higher chance of producing a fertilized first cocoon. (VII) A trade-off existed between producing many small versus few large young.     

Life history of the oligochaete Enchytraeus coronatus (Annelida, Enchytraeidae) in agar culture

Abstract. Life-history attributes and the reproductive biology of Enchytraeus coronatus (Oligochaeta, Enchytraeidae) were studied in agar cultures, as a basis for using this species in toxicity tests and other experimental studies. The use of agar allows daily examination of the behavior, reproduction, and mortality of the worms. At 20°C, E. coronatus had a generation time of 21–22 days and an embryonic period of 7–10 days. Hatch rate was high (>80%) and worms became sexually mature 8–10 days after hatching. Cocoon production and number of eggs per cocoon were positively correlated. Mortality in the first stages of the life cycle (embryonic period and first week after hatching) was about 38%. Three different periods of cocoon production were distinguished: a period characterized by a regular increase in the number of cocoons produced (weeks 4–9), a period of production at a constant high rate of ∼5 cocoons per week (weeks 10–20), and a period of decline in production rate (weeks 21–32), down to a mean value of 1.6 cocoons per adult in the last week. This study showed that synchronized agar cultures of cohorts can provide adults for long-term experimental tests, with the age for optimum reproduction being 8–20 weeks.     

Reproductive potential of the earthworm Eisenia foetida

Summary Regression equations are provided for the earthworm Eisenia foetida with respect to age at which 50% of the population became clitellate at 25° C in relation to population density in activated sludge and in horse manure. Regression equations are provided for progeny per cocoon versus weight of cocoon, and weight of cocoon in relation to weight of parent; from these an equation is derived for progeny per cocoon relative to worm weight. Regression equations are given on (a) number of cocoons produced per adult in relation to age and population density from onset of adulthood to median peak production of cocoons, age 10 weeks, and from age 10 weeks to age 27 weeks, and (b) weight of worm in relation to population density and age between ages 5 and 27 weeks. From (a) and (b) a family of equations (c) are derived giving progeny per cocoon in relation to age of adult and population density. From equations (a) and (c) two families of equations are generated giving progeny per adult in relation to ascent to, and descent from, the median week of peak cocoon production in relation to population density. Data also are provided on age at which reproduction terminates in relation to population density, optimum population density for reproduction, and hatchability.     

Predicting efficiency of selection on reproductive traits in the medicinal leech Hirudo medicinalis L

A set of selection measures for increasing reproduction efficiency in Hirudo medicinalis has been developed. The optimal values of reproductive traits corresponding to the highest progeny number were determined and recommended. The probability of correlated selection response in traits "number of threads in a cocoon" and "weight of threads" was estimated. Based on earlier results on phenotypic variation and heritability of reproductive traits in medicinal leech, efficiency of different selection modes was predicted.     

Studies on the Relationships Between Earthworms and Soil Fertility

The rate of cocoon production in British Lumbricidae differs considerably from species to species and is greatly affected by soil temperature, moisture and the food supply of the adult worms. Only two out of fourteen species studied frequently produced more than one worm per cocoon, and twins were only recorded once in many hundreds of observations on four species of the genus Lumbricus.
The incubation period of the cocoons and the growth period to sexual maturity varies from species to species and also depends on the time of year at which the cocoons are deposited and the young worms emerge.     

体重、年龄和应激刺激对菲牛蛭繁殖性能的影响

采用单因素方差分析等方法探讨人工养殖条件下体重、年龄和环境应激对菲牛蛭繁殖性能的影响。结果表明,体重对卵茧大小和湿质量及幼蛭体长、体宽和湿体质量无显著影响(P>0.05),而亲蛭性腺系数、平均产茧数、孵化率和平均孵化数在各组间差异显著(P<0.05),其中体重10～15g组明显高于其他组(P<0.05);年龄对亲蛭平均产茧数、孵化率、平均孵化数和卵茧大小影响显著(P<0.05),其中2～3年龄亲本具有最高的平均产茧数、孵化率和平均孵化数,8月龄亲本所产卵茧个体明显大于其他年龄组(P<0.05),而卵茧湿质量及幼蛭体长、体宽和湿体质量在各年龄组间均无显著差异(P>0.05)。环境应激对亲蛭的平均产茧数、孵化率和平均孵化数影响显著(P<0.01),其中1～2次操作应激下的平均产茧数和孵化数最高,无操作刺激下的孵化率略高于1～2次操作刺激(P>0.05),而卵茧大小和湿质量及幼蛭体长、体宽和湿体质量在各应激处理组间无显著差异(P>0.05)。本研究获得了菲牛蛭繁殖的最佳体重和年龄,建议繁育生产中挑选10～15g和2～3年龄的个体作为亲本,期间进行1～2次操作刺激可提高其繁殖性能。     

The cocoon and hatchling of Nais variabilis (Naididae: Oligochaeta)

SUMMARY. 1. The cocoon of Nais variabilis is ovoid with an average length of 527.6±8.4 μm (95% CL) and an average breadth of 378.1±9.9 μm. The cocoon wall is about 7 μm thick and the average volume of the cocoon cavity is 0.035±0.002 mm³. Most cocoons contain one ovum; only single-ovum cocoons developed.
2. Cocoons hatch in about 80 days at room temperature (average 20°C. range 17–23°C). Hatchlings emerged posterior end first and were, on hatching, about one-third the adult live length of 8.5±10.3 mm.
3. It is estimated that adult Nais variabilis are unlikely to produce more than an average of ten cocoons each. Adults ceased asexual budding on becoming nature and died shortly after the completion of cocoon production. There was a brief co-existence of generations.     

Antagonistic effects of energy status on meal size and egg-batch size of Aedes aegypti (Diptera: Culicidae).

The effect of sugar feeding on egg-batch size in Aedes aegypti was examined in a way that would distinguish between the roles of a recent sugar meal (full vs. empty crop) and of repeated sugar feeding and digestion (high vs. low energy reserves). Egg numbers of females representing the four combinations of these characteristics (full crop/high reserves, full crop/low reserves, empty crop/high reserves, empty crop/low reserves) were counted during their first gonotrophic cycle. In addition, the sizes of their replete sugar meals and human blood meals were measured to understand the interactions between them. Results demonstrated that blood-fed mosquitoes with full crops and low energy reserves produced the fewest eggs (mean = 56.2); those with empty crops and high energy reserves produced the most eggs (mean = 84.6); and those with the opposite combinations (full/high and empty/ low) had an intermediate fecundity (mean = 75.2 and 76.9, respectively). This ranking of fecundities did not correspond to blood-meal size ranks, owing to direct and indirect effects of energy reserves on meal sizes and egg number. Full-crop females with low reserves ingested the smallest blood meals (mean = 0.62 mg) and had the lowest fecundities. Full-crop females with high reserves ingested more blood (mean = 0.82 mg) and produced more eggs. But empty-crop females with low reserves ingested the largest blood meals (mean = 1.19 mg), yet produced significantly fewer eggs than their high-reserve counterparts, which took smaller blood meals (mean = 0.99 mg). These results demonstrate extremes in the reproductive penalty of crop sugar and the reproductive reward of digested sugar. Energy reserves and an empty crop are similarly valuable in promoting fecundity at the time blood is taken.     

Reproductive ecology of Dendrocoelum lacteum (Turbellaria) in a rapid stream in southern Sweden and comparisons with a lake population

Rapid streams in southern Sweden are often inhabited by the triclad Dendrocoelum lacteum. Outside Fennoscandia, this species is mainly restricted to lentic habitats. The normal food refuge for D. lacteum, the isopod Asellus aquaticus, is seldom found in the stream habitats, and probably the prey here is the amphipod Gammarus pulex. With respect to spatial and trophic niche components, it seems that D. lacteum has taken the place of Crenobia alpina in southern Sweden.Two parameters of reproduction, namely production of cocoons and of hatchlings, both peaked approximately one month later in a stream than in an adjacent lake. This difference was attributed to a low temperature regime in the stream. Number of hatchlings per cocoon and cocoon sterility were higher in the stream than in the lake. Similar cocoon and hatchling output per adult was found in the two habitats, indicating a similar reproductive effort for the two triclad populations. I suggest that D. lacteum, by virtue of its choice of microhabitat, viz., under stones, is not so vulnerable to the more severe stream environment.     

Infectivity of Neoaplectana carpocapsae and Heterorhabditis heliothidis to pupae of the parasite Apanteles militaris

The infectivity of Neoaplectana carpocapsae and Heterorhabditis heliothidis to Apanteles militaris, a gregarious parasite of the armyworm, was deterntined at 100. 1,000, 5,000, and 10,000 nematodes per petri dish. For both nematode species, the percentage of infected A. militaris within a cocoon cluster decreased as inoculum levels decreased. At the highest inoculum level, N. carpocapsae infected an average of 32% of the parasite pupae within a cocoon cluster, whereas H. heliothidis infected an average of 22%. Covariance analysis indicated, however, that N. carpocapsae had significantly greater infectivity than did H. heliothidis. Some of the dauer juveniles of N. carpocapsae on the body of the armyworm contacted the emerging parasites and eventually became enveloped within the silken cocoons. Dauer juveniles produced by N. carpocapsae in parasite pupae could not penetrate and escape from silken cocoons even when the cocoons were placed in a moist environment.     

Breeding system ofDaphniopsis ephemeralis: adaptations to a transient environment

Populations of the vernal cladoceranDaphniopsis ephemeralis are found in woodland ponds throughout southern Ontario. The species reproduces by cyclic parthenogenesis, and genotype frequencies at allozyme loci are ordinarily in good agreement with Hardy-Weinberg expectations. Occasional heterozygote deficiencies are apparently the consequence of admixture of ephippial hatchlings produced in temporally separated bouts of sexual reproduction. Considerable heterogeneity in genotypic frequencies exists among local populations in southwestern Ontario, indicating that gene flow among populations is restricted. Inbreeding coefficients suggest that populations receive an average of 0.3 migrants per generation. The completion of a sexual life cycle is made possible despite the brief persistence of populations by the emergence of males from ephippial eggs and by the production of equal numbers of male and female progeny in the first parthenogenetic brood.     

A comparative survey of the Erpobdella octoculata (L.) populations in the Afon Crafnant,N. Wales,above and below an input of zinc from mine-waste

The life-cycle, growth, and reproductive success of a stream-dwelling Erpobdella octoculata (L.) were studied over a 19-month period at two sites, one above and one below an input of zinc pollution from mine-waste. Samples were collected monthly using quadrats. The blotted wet weight of all leeches was measured. An histological technique was employed to determine sexual maturity. Reproductive success was measured by estimating the numbers of cocoons and number of eggs per cocoon at each site. A wet ashing technique, followed by analysis using flame atomic absorption spectrometry, was used to determine the extent of bioaccumulation of zinc. Total zinc and other metals in the waters were similarly analysed.Differences were found between the populations at the two sites. The life-history of the Erpobdella above the contamination was comparable to that found for other stream-dwelling populations previously studied. Observed differences in densities of leeches between sites in the present and in previous studies by other workers may be due to recorded differences in the abundance of available food. At the polluted site, there was evidence that (a) there was a delay in cocoon deposition and hence hatching of young, (b) the adult leeches produced more misshapened and empty cocoons, (c) no relationship existed between the area of cocoons with eggs and number of eggs present, and (d) the proportion of young to adult leech was smaller at the polluted site. At the polluted site, higher levels of zinc were recorded in the tissues of Erpobdella and there was some evidence for the active intake of zinc by the leech.     

Life cycle of the earthworm Octodrilus complanatus (Oligochaeta, Lumbricidae)

The earthworm Octodrilus complanatus (Dugès, 1828) is a large-sized species often found in grasslands and pastures, with a wide distribution area in Europe and North Africa. In order to determine the characteristics of its life cycle, earthworms of this species were cultured from hatching until they reached maturity. Development rates, cocoon production, and incubation period were recorded under controlled conditions. On average, individuals of O. complanatus reared in isolation from hatching reached maturity at day 150 and body weight at maturity ranged between 6 and 8 g. The cocoon incubation period was 66 days, and only one individual hatched from each cocoon, with a mean hatchability of 55%. Reproductive rate showed by O. complanatus after mating was of 52 cocoons per earthworm per year. The results showed that the life cycle of O. complanatus was characterized by both low growth rates and low fecundity. Specific features in life history traits of O. complanatus were compared with those of species of the genus Octodrilus and other related earthworms.     

Laboratory studies on the life-histories of four enchytraeid worms (Oligochaeta) which inhabit sewage percolating filters

The life-histories of four enchytraeid worms, Lumbricillus rivalis, Enchy-traeus coronatus, E. buchholzi, and E. albidus which occur in sewage percolating filters, were studied under laboratory conditions at 8 , 15 and 20°C. The number of ova per cocoon varied from 0 to 50 (L. rivalis), 0 to 33 (E. coronatus), 1 to 9 (E. buchholzi) and 0 to 22 (E. albidus). The mean number of ova per cocoon was highest at 15°C for all species except E. coronatus which had a highest mean value at 8°C. The number of ova in cocoons was correlated with cocoon length (P < 0.001) for all species. Cocoon production usually increased with temperature ranging from 0.8 cocoons per adult per week at 8°C to 2.0 at 20°C for L. rivalis, and from 1–4 to about 2.6 for E. coronatus and E. buchholzi. The total number of ova produced by each E. coronatus (350 at 8°C to 550 at 20°C) was similar to that produced by each L. rivalis (600 at 8°C to 350 at 20°C) and was about five times greater than the total numbers produced by the other two species. Cocoon and ova production and the number of ova per cocoon varied with the age of the adult, usually reaching a peak soon after maturity. Hatching success was low and generally 40–50 % of ova failed to develop; subsequent mortality among immature worms was about 10–20%. Growth was more rapid at the higher temperatures; L. rivalis matured in about 26 days at 20°C, the clitellum forming when the worm was 13–14 mm long; data for the other species are 13 days and 5–6 mm (E. coronatus); 16 days and 3–4 mm (E. buchholzi); 28 days and 13–14 mm (E. albidus). The maturation period at 8°C was at least twice that at 20°C. The generation period (cocoon to cocoon) was about a month at 20°C for all species except E. albidus (2 months), but as some species had longer reproductive periods than others the actual number of generations per year was highest in E. buchholzi, 7.0 per year, and lowest in E. albidus, about 3.3 per year, At 8°C all four species had between 1.4 and 2.8 generations a year. A comparison of expected and observed population densities of L. rivalis and E. coronatus in a sewage percolating filter showed that neither achieved values approaching their potential summer densities although ample food was apparently available. Of the four species studied only E. buchholzi produced viable ova without pairing.     

Multiple mating increases cocoon hatching success in the earthworm Eisenia andrei (Oligochaeta: Lumbricidae)

In simultaneous hermaphrodites with reciprocal mating, multiple mating may be a male strategy that conflicts with female interests, and therefore an intra‐individual sexual conflict regarding the number of matings may be expected. The evolutionary outcome of this sexual conflict will depend on the costs and benefits that extra mating entails for each sexual function. In the present study, we investigated the costs and benefits of multiple mating on cocoon number, cocoon mass, and cocoon hatching success in the redworm Eisenia andrei, a simultaneous hermaphrodite with reciprocal insemination, by manipulating the number of matings with different partners. We did not detect any reduction in the female reproductive output (number and mass of cocoons) with increasing number of mating partners. However, we found that multiple mating showed benefits for female reproduction that increased the hatching success of the cocoons. This effect may be a result of increased quantity and/or diversity of sperm in the spermathecae of multiple mated earthworms. Further studies are required to clarify the mechanism underlying the increased cocoon hatching success when redworms engage in multiple matings. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Seasonal population dynamics of Eisenia fetida (Savigny, 1826) (Oligochaeta, Lumbricidae) in the field

In order to assess the response of epigeic earthworms to seasonal changes we monitored the population dynamics of Eisenia fetida (Oligochaeta, Lumbricidae) in a manure heap in the field during a year. Earthworms were hand-sorted from five 0.25 x 0.25 x 0.20 m blocks around the heap in November (autumn) 1999 and in January (winter), April (spring) and August (summer) 2000 to determine earthworm population dynamics. Earthworms of each block were classified into different age classes: mature, preclitellate, juvenile, hatchling and cocoon, and afterwards counted and weighed. Seasonality had a strong effect on the density, biomass and reproductive activity of the population. The population of E. fetida was characterized by a high density of individuals and the predominance of mature individuals throughout the year. Maximum density, mating activity and size of cocoons were achieved in spring, but there were not changes in the number of cocoons per mature earthworm throughout the year. Unexpectedly, the smallest cocoons were produced in winter by the largest individuals. These results suggest that E. fetida is able to allocate resources to growth and/or reproduction in response to environmental fluctuations.