Multiple paternity and mate competition in non-selfing,monogamous, egg-trading hermaphrodites

In animals in which the two sexes invest relatively similar amounts of resources in their young, the number of mates is expected to affect male and female reproductive success similarly and gender conflicts on the number of mates may not arise. Correspondingly, in non-selfing, simultaneous hermaphrodites with long-term monogamy, the two partners are expected to alternate repeatedly their sexual roles and invest similarly in their offspring. Therefore, the gender conflict on the number of mating partners should not arise. However, when >2 conspecifics are present, hermaphrodites are known to plastically adjust their behavior and sex allocation and compete for mating repeatedly in the male role. We tested whether this leads to multiple paternities of single egg clutches in experimental replicates of small and large groups of non-selfing, egg-trading, behaviorally monogamous polychaete worms (Ophryotrocha diadema) by using neutral genetic markers to estimate paternity. Multiply fertilized egg cocoons were common in these worms; two or more individuals succeeded in fertilizing the same egg cocoon and mate competition increased with group size. Multiply fertilized egg cocoons had a higher proportion of eggs developing into mature worms than singly fertilized egg cocoons. Possibly singly fertilized cocoons had a lower fertilization rate owing to low sperm counts and aflagellate sperm.     

A comparison of reproduction, growth and acute toxicity in two populations of Tubifex tubifex (Müller,1774) from the North American Great Lakes and Northern Spain

Reproduction in Tubifex tubifex is being used as part of a suite of indicators of sediment toxicity in Canada and Spain, and reproduction of T. tubifex is being considered as a component of sediment objectives for environmental regulation and clean-up in the Canadian Great Lakes. The data being used to set these reproductive targets have been developed from a single culture of T. tubifex from Lake Erie. The plasticity of this particular species is well known and before it can be adopted widely as a test organism it is necessary to determine whether a single culture source should be used or if cultures derived from different populations respond similarly. A series of experiments with two cultures, one from Lake Erie the second from a small mountain stream in Northern Spain have shown that the Spanish worms appear to produce fewer cocoons per adult (mean 8.6 S.D. 1.0) than those from Lake Erie (mean 10.4 S.D. 0.3) at 22.5 °C, a standard test temperature. The number of young produced per adult by the Spanish culture is also lower (mean 19.0 S.D. 4.6) than the L. Erie population (mean 30.6 S.D. 2.3), however, the Spanish population has higher reproductions rates at a lower temperature. The Spanish worms also have lower and more variable growth rates than the Canadian population. There also appear to be slight differences in the sensitivities to toxicants, with the Canadian worms having higher LC50s for copper, chromium and cadmium. While there are differences in the responses in the two cultures these are not considered to be sufficient to invalidate the use of either population in a standard bioassay protocol as long as appropriate calibration and validation are undertaken.     

Reproductive ecology of Naididae (Oligochaeta)

Asexual reproduction is employed by species of Naididae during favorable environmental conditions. In species characteristic of aquatic habitats subject to rapid fluctuations in water levels and temperatures most individuals in a population become sexually mature, and there is degeneration of the alimentary tract, a shortened breeding season, and aclitellar formation of cocoons. Cocoon deposition allows a population to survive periods of environmental stress. Sexual reproduction predominates near the edges of ranges where stress conditions are more prevalent. Species that reproduce sexually in spring are intolerant of summer conditions, while those that breed in autumn are intolerant of winter. Species typical of environmentally stable bodies of water usually have a minority of the population mature at any time, an extended breeding season, and the ability to continue feeding while mature; thus, adaptations for survival in unstable habitats are lacking.     

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.     

A review of the biology of British Naididae (Oligochaeta) with emphasis on the lotic environment

SUMMARY. Naidids live in a wide range of aquatic habitats but are particularly important numerically as part of the benthic fauna of rivers with stony substrates. In general they graze on bacteria and algae although some, particularly Chaetogaster spp., are mainly predaceous, and C. limnaei vaghini is a parasite of molluscs, chiefly Gastropoda. Food selection seems to be based largely on particle size although the food quality of the particles within the appropriate size-range influences rates of growth and reproduction. Major factors determining the distribution and abundance of naidid species are the nature of the substratum and the presence and kind of vegetation. Plants with a highly dissected form, a thick growth habit, and which permit the greatest periphyton development generally support the most abundant naidid populations. The oligochaete fauna of coarse substrates (stones and gravels) is often dominated by the Naididae but both species-richness and abundance of naidid populations are generally reduced where fine substrates (silts and muds) occur. The occurrence of worms within the substratum is also determined by its nature; naidids penetrate to depths of 20–70 cm in coarse substrates but rarely penetrate below 6 cm in mud. The principal factor limiting both depth penetration and the dominance of naidids in fine substrates is probably oxygen availability. Naidids reproduce both asexually and sexually, the former method predominating for most of the year. Asexual reproduction usually involves the budding-off of zooids but a few species fragment. Sexual reproduction is often infrequent; populations of some species produce few or no sexually mature individuals. In mature worms asexual reproduction virtually ceases. In populations that produce mature individuals there is apparently one sexual generation a year, usually occurring during the summer and autumn. Adults die soon after laying their cocoons. In general, naidids are most abundant during the summer months when rates of growth and asexual reproduction are stimulated by higher temperatures and a plentiful supply of food. A few species, however, e.g. Nais elinguis and Paranais litoralis, are more abundant in the spring. The response of naidid species to different kinds of pollution is varied but generally organic enrichment of rivers which have stony substrates results in a considerable (ten- to twenty-fold) increase in naidid abundance. Nais elinguis, N. barbata, N. communis, N. variabilis, and Chaetogaster diaphanus are often abundant in such rivers, the foremost species reaching densities of 200 000 m^?2. Nais alpina, N. bretscheri, and N. pardalis appear to be relatively intolerant of organic enrichment. A substantial increase in naidid abundance is also promoted by the deposition of biologically inert particles (coal dust, china clay, sand) on the river bed. The importance of Naididae in relation to pollution surveillance of fresh waters is discussed.     

Toxicity of Santander Bay Sediments to the Euryhaline Freshwater Oligochaete Limnodrilus hoffmeisteri

The freshwater euryhaline oligochaete Limnodrilus hoffmeisteri was selected for use in bioassays with polluted sediment from Santander Bay. It is easy to culture; is tolerant of low to moderate, up to 15‰, salinity; and is common in oligohaline conditions in European and North American estuaries. Worms were collected from an estuarine population and kept in unpolluted sediment for between 2 and 4 weeks, under laboratory conditions, at 7–8.5‰ salinity and 22.5 °C. Sediment from different sites in Santander Bay were sieved through 250 μm mesh and adjusted to a salinity of approx. 7‰ prior to the bioassays, either by adding sea water or dechlorinated tap water to the overlying water. High levels of ammonia in some sediment, which may confound results in ecotoxicity bioassays, were reduced by oxidation of the sediments in shallow trays. Sediment bioassays were performed with sexually mature Limnodrilus hoffmeisteri worms in 250-ml beakers, with a 1:3 ratio of sediment:water and 4 worms per baker. Endpoints in the 14-day bioassay were % mortality, adult final biomass, % adults that have shown resorption of the clitellum, number of cocoons, and burrowing behaviour. It was possible to rank the sites according to their toxicity using both mortality rates and sublethal effects. The control site had the following values for the endpoints: 5% mortality, ( ) 2.40 ± 1.52 cocoons per beaker and 1.271 ± 0.470 mg dw adult final biomass. The most toxic sediment resulted in 65% mortality, resorption of the clitellum in 67% of the adults, no production of cocoons and a low final biomass ( =0.681 ± 0.489 mg dw per adult). A second site had high mortality (60%) and no reproduction, although resorption of the clitellum did not occur in surviving animals. The remaining sites showed similar mortality (35–42%), and at only one of them was low reproduction observed (0.8 ± 0.447 cocoons per beaker). Behavioural effects, measured as length of galleries in a fixed area of the test-vessel at the end of the bioassay, were significant compared with control at only one site. Multivariate analysis showed the mortality gradient to be the strongest, with a second unassociated gradient representing clitellum resorption. The mortality gradient was associated with Cu and Zn concentration, and PAH and Pb possibly with resorption.     

Unique mating behavior,and reproductive biology of a simultaneous hermaphroditic marine flatworm (Platyhelminthes,Tricladida, Maricola)

Flatworms generally are simultaneous hermaphrodites that exhibit various kinds of mating behavior. Here we report on the mating behavior and reproductive biology of the planarian Paucumara falcata. We recognized three phases in its mating behavior: a courtship, copulation, and postcopulatory phase. During the last‐mentioned phase, the partners showed a unique and very characteristic behavior in which their bodies intertwined, forming a spiral. Histological study of partners in copula revealed that the sclerotic tip of the musculo‐parenchymatic organ pierces the body wall of the partner and then becomes lodged in its parenchyma, suggesting that this organ may act as an anchor, thus stabilizing the worms during copulation. Similar organs in other species of marine triclad may also perform a stabilizing role during copulation. During copulation in individuals of P. falcata, sperm transfer was reciprocal or only unilateral. Copulation duration ranged 13–35 min (average 20 ± 5 min), irrespective of whether the mating was successful (i.e., resulted in the production of fertile cocoons). The spiraling phase lasted on average 10 min; some worms did not show the postcopulatory spiraling phase during their mating behavior. After successful copulation, an individual worm produced 1–12 fertile cocoons over a period of 1–17 days; from a cocoon hatched either one young (in 70% of the cases), or two young worms.     

Reproduction of Lumbricillus rivalis (Levinsen) in laboratory cultures and in decaying seaweed

The cocoon production of 144 Lumbricillus rivalis cultured in pairs at 10 ± 1 °C was high over the first 2 weeks of breeding activity and then declined, chiefly because of high mortality. Cocoon deposition lasted for between 1 and 16 weeks, eight pairs of worms producing cocoons for 9 weeks and one pair for 16 weeks. During the total period of cocoon deposition over 9000 eggs (mean 17·4 per cocoon) were deposited. Two decaying wrack bed populations of L. rivalis showed a low level of cocoon and egg production in autumn, rising to an annual maximum in late winter/early spring. In these populations the mean egg content varied seasonally from 17·1 to 47·8 eggs per cocoon. When cocoons in the laboratory were transferred from the site of deposition to incubation dishes 31% hatched, but those left in the substrate showed a 92% hatch. In the naturally occurring populations 19% of the cocoons detached from seaweed fronds hatched, but 62% of those left in situ. Eggs and worm embryos developed to relatively late stages in most cocoons, whatever the rate of hatching; development often continued for up to 2 months after deposition without hatching. Over 50% of the fertile eggs in cocoons from decaying wrack hatched and developed to 5 mm worms.     

The population biology ofPlanaria torva (Müller) (Turbellaria,Tricladida)

Summary 1. A quantitative study of the population biology ofPlanaria torva (Müller) living in a productive lake, based on monthly samples over a period of 20 months, is presented. 2. Samples of triclads, their cocoons and other organisms were taken from bricks placed at a shallow depth in the lake. They provided relative information on changes in population size structure and demonstrated the temporal breeding of this triclad. 3. Cocoon production byP. torva began in January and reached a peak in March–April. This was followed by a peak in recently-hatched young during May–June. Adults reached a maximal proportion of the total population in January and most had died by July. 4. Changes in size structure suggest that intra-specific competition occurred during June–July following the recruitment of young. This hypothesis is supported by other independent evidence. 5. Attempts to measure fecundity based on cocoons and young gave very different results attributed to the sampling technique.     

Growth,reproductive biology and life cycle of the vermicomposting earthworm, Perionyx ceylanensis Mich. (Oligochaeta: Megascolecidae)

In the present study, an attempt has been made to study the growth, reproduction and life cycle of the earthworm, Perionyx ceylanensis Mich. in cowdung for the period of 340 days. Results showed that the overall mean growth rate was 1.79, 1.57 and 1.34 mg/worm/day respectively for the worms cultured singly, in batches of four and eight. Cocoon production rate was found between 0.85 and 0.94 cocoons/worm/day and the hatching success between 74.67% and 82.67%. The majority of the cocoons (95.16–96.77%) hatched only one hatchling. Worms raised singly also produced viable cocoons indicating that P. ceylanensis reproduce parthenogenetically. The life cycle of the worms cultured singly was ±57 days and it was ±50 days for the worms cultured in batches of four and eight. There is a vast scope to utilize P. ceylanensis for vermiculture practices due to short period of life cycle.     

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.     

Biological studies upon Enchytraeus variatus Bouguenec & Giani 1987 in breeding cultures

The life history of an enchytraeid worm, Enchytraeus variatus, was studied under laboratory conditions at 18–22 °C. This species can reproduce simultaneously by asexual (architomy) or sexual reproduction. The number of ova per cocoon varies from 5 to 20 (x = 10.9). The generation period (from cocoon to next cocoon) varies from 14 to 39 days (x = 26.1) according to the period of the year. The number of generations per year is between 7.3 and 26.1 (x = 14). A mature worm can lay between 23.7 and 25.8 cocoons during its life (254 days as maximum observed) at a mean rate of 0.12 cocoon worm^–1 day^–1. Experimental cultures were carried out to determine the structure, density and biomass of the populations. A maximal density of 1 396 314 worms was recorded after 85 days of culture. Net production reached 21.48 g m^–2 day^–1 after 26 days in a culture initiated from cocoons.     

The structure and distribution of external sense organs in newly hatched and mature earthworms

Fully mature adult Eisenia foetida sensory buds are abundant on the prostomium and the first segment. In subsequent segments they are restricted to the anterior half where they form a single row aligned with the setae and encircling the worm. In the more posterior regions of the worm the buds are widely separated and fewer. The surface of each bud is a raised circular or oval area from which 15 to 100 so-called sensory hairs arise, being cylindrical and apparently flexible. The number of these projections decreases toward the posterior end of the worm. In worms newly emerged from egg cocoons, the general pattern of distribution and external form of sensory buds resembles that of adults, but the buds are much fewer and smaller than in adults. Although these worms emerge with their definitive adult number of segments, new buds and additional sensory projections are formed during post hatching development.     

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.     

Switch from Asexual to Sexual Reproduction in the Planarian Dugesia ryukyuensis

Many metazoans convert the reproductive modes presumably depending upon the environmental conditions and/or the phase of life cycle, but the mechanisms underlying the switching from asexual to sexual reproduction, and vice versa, remain unknown. We established an experimental system, using an integrative biology approach, to analyze the mechanism in the planarian, Dugesia ryukyuensis (Kobayashi et al., 1999). Worms of exclusively asexual clone (OH strain) of the species gradually develop ovaries, testes and other sexual organs, then copulate and eventually lay cocoons filled with fertilized eggs, if they are fed with sexually mature worms of Bdellocephala brunnea (an exclusively oviparous species). This suggests the existence of a sexualizing substance(s) in sexually mature worms. Random inbreeding of experimentally sexualized worms (acquired sexuals) produces an F1 population of spontaneous sexuals (innate sexuals) and asexuals in a ratio of approximately 2:1. All regenerants from various portions of innate sexuals become sexuals. In the case of acquired sexuals, head fragments without sexual organs regenerated into asexuals though regenerants from other portions became sexuals. Thus, we conclude that neoblasts, the totipotent stem cells in the planarians, of acquired sexuals remain "asexual" and the worms require external supply of a sexualizing substance for the differentiation of sexual organs and gametes. On the other hand, some, if not all, neoblasts in innate sexuals are somehow "sexual" and do not require external supply of a sexualizing substance for the eventual differentiation of themselves and/or other neoblasts into sexual organs and gametes. It is also shown that sexuality in acquired sexuals is maintained by the putative sexualizing substance(s) of their own. The sexualization is closely coupled with cessation of fission, and the worms seem to have an unknown way of controlling the karyotype. Our integrative approach integrates multiple fields of study, including classic breeding, regeneration, and genetics experiments, as well as karyotyping, and biochemical and molecular biological analyses; none of which would have revealed much about the intricate mechanisms that regulate sex and fission in these animals.     

Cold hardiness and geographic distribution of earthworms (Oligochaeta,Lumbricidae, Moniligastridae)

Cold hardiness of 12 species and 2 subspecies of earthworms from Northern Eurasia was studied. Supercooling temperatures, the water content and the thresholds of tolerated temperatures of worms and their cocoons were determined. The threshold values varied within ?1…?35°C for worms and within ?1…?196°C for cocoons. Earthworms of 4 species and 2 subspecies survived freezing. Cocoons of all species except Eisenia fetida possessed a protective dehydration mechanism which prevented their freezing. During wintering at subzero temperatures, earthworms lost up to 20% of water, cocoons up to 37%. Species of the same life form can overwinter at different phases and have different cold hardiness values. On the whole, epigeic and epi-endogeic species (except for Eisenia fetida) were more resistant to cold than endogeic ones. The following preliminary classification of earthworms according to their tolerance to negative temperatures is proposed: (1) both onthogenetic phases are tolerant; (2) only cocoons are tolerant; (3) both onthogenetic phases are intolerant. The geographic distribution of all the studied species (except for Eisenia nordenskioldi nordenskioldi) is partially or completely limited by insufficient resistance of the worms to negative temperatures. A significant cold hardiness of cocoons of most species is nonadaptive, since the worms hatched from the eggs in spring die without having enough time to reach maturity and to lay cocoons before the onset of subzero temperatures. Only 3 species (Eisenia nordenskioldi nordenskioldi, Eisenia atlavinyteae, and Dendrobaena octaedra) can live in permafrost regions; this is the main reason for a drastically reduced diversity of earthworm assemblages in eastern Siberia except for its southern, mountain parts. In general, the reasons for the impoverishment lie in the modern climatic conditions correlated with the ecophysiological capacities of earthworms.     

A mechanism for self-inhibition of population growth in the flatworm Mesostoma ehrenbergii (Focke)

Summary By producing subitaneous eggs the rhabdocoel Mesostoma ehrenbergii may quickly increase the population size. The production of this kind of eggs by young worms is inhibited by the presence of adults in the culture medium. The inhibition is specific, and it is observed also if young and adult worms are not in direct contact, or if young worms are added to the water where adults had been previously kept: there is therefore evidence for a chemical mediator. The sensitivity to the inhibition occurs during the first five days after birth, with a maximum on the 2nd to 3rd day.     

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.     

Switching from asexual to sexual reproduction in the planarian Dugesia ryukyuensis: change of the fissiparous capacity along with the sexualizing process

Asexual worms of fissiparous strain of the planarian Dugesia ryukyuensis switch from asexual to sexual reproduction, if they are fed with sexually mature worms of Bdellocephala brunnea. This suggests that the sexually mature worms have a sexualizing substance(s) that induces the sexuality in the asexual worms. Here, we found by analysis of the sexualization that the cessation of the fission, namely their asexual reproduction, occurs immediately after the acquisition of sexuality. This result suggests that the downstream mechanisms induced by the putative sexualizing substance in B. brunnea become responsible for the cessation of fission. We also found that the decapitation triggers fission in the worms even after the acquisition of sexuality if they are not sexually mature, while the fully sexualized worms never fission even though they are decapitated. This result suggests that the cessation of fission takes place via at least two steps: (1) the mechanisms associated with the cephalic system; (2) other mechanisms independent of cephalic control.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.