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.     

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.     

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.     

Seasonal dynamics and emergence pattern of a tropical earthworm,Drawida calebi (Oligochaeta)

Cocoon morphology (shape, size, colour and ornamentation), hatching, and seasonal dynamics of cocoon and juveniles of D. calebi have been studied in an Indian pasture site. The average live weight of cocoon is 15.17 mg, length/diameter 0.94, and moisture content 72.5 g%. Just-hatched juveniles weigh on an average 10.4 mg, and are 10.5–17.5 mm in length. The incubation period lasts 56.33 ± 2 days during November—December in laboratory cultures (25°C, and moisture 15 g%). A maximum of 64.0 live cocoons/m² with a live weight of 0.971 g/m² were deposited during October—November 1977, but very few during summer months. A surface-living species, D. calebi shows single peak emergence pattern in December—January. It is estimated that 78 cocoons (live and dead)/m² (780,000/hectare), were produced in the field by D. calebi population in a year (1976–77). Thus the total cocoon production is 11.83 kg/hectare/year. It is estimated that 3 cocoons/individual/year are produced by D. calebi in field conditions.     

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.     

Phenology and nutrition of miombo woodland trees in Zambia

Seasonality in nitrogen (N) and phosphorus (P) concentration in soil and shoots of five Brachystegia-Julbernardia (miombo) woodland trees was studied from September 1991 to March 1993 at two regrowth miombo sites in central Zambia. Shoot growth started in the dry season (September–November) and lasted until April during the 1991/92 season but had virtually ceased by January 1993 during the 1992/93 season. The shoot growing season was associated with low foliar N/P ratios. These ratios were much lower (<5) during the 1991/92 season than in the 1992/93 season (12–15). The increase in foliar N/P ratios after the shoot growing season was caused by a sharp drop in foliar P concentration, apparently due to reabsorption before leaf fall. There were no annual variations in biomass N concentration in contrast to P. During the 1992/93 growing season P concentrations in foliage and wood were a quarter and a third, respectively, of the 1991/ 92 levels. Since the short shoot growing season observed during the 1992/93 season is typical of savanna woodland trees in southern Africa, the high biomass P concentration and longer growing season in 1991/92 season were exceptional and may have been related to reduced competition by shallow rooting herbaceous plants caused by the severe drought of that season.     

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, ?? , ??–??.     

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。     

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.     

Behavioral manipulation of host caterpillars by the primary parasitoid wasp Cotesia glomerata (L.) to construct defensive webs against hyperparasitism

Many parasitoids control the behavior of their hosts to achieve more preferable conditions. Decreasing predation pressure is a main aim of host manipulation. Some parasitoids control host behavior to escape from their enemies, whereas others manipulate hosts into constructing defensive structures as barriers against hyperparasitism. Larvae of the parasitoid wasp Cotesia glomerata form cocoon clusters after egression from the parasitized host caterpillar of the butterfly Pieris brassicae. After the egression of parasitoids, the perforated host caterpillar lives for a short period and constructs a silk web that covers the cocoon cluster. We examined whether these silk webs protect C. glomerata cocoons against the hyperparasitoid wasp Trichomalopsis apanteroctena. In cocoon clusters that were not covered by silk webs (bare clusters), only cocoons hidden beneath others avoided hyperparasitism. In covered cocoon clusters, both cocoons hidden beneath others and those with a space between them and the silk web avoided hyperparasitism, whereas cocoons that contacted the silk webs were parasitized. The frequency of cocoons that were hidden beneath others increased with the increasing number of cocoons in a cluster, but the defensive effect of cluster size was thought to be lower than that of silk webs. However, the rate of hyperparasitism did not differ between covered and bare clusters when we allowed the hyperparasitoids to attack the cocoon clusters in an experimental arena. This result was thought to have been caused by low oviposition frequency by these hyperparasitoids. As a result, silk webs did not guard the cocoons from hyperparasitoids in our experiments, but would protect cocoons under high hyperparasitism pressure by forming a space through which the ovipositors could not reach the cocoons.     

The role of spider cocoons in controlling desiccation

Summary The abilities of the cocoons of the spiders Mecynogea lemniscata and Argiope aurantia to protect the enclosed egg and spiderling stages from desiccation were investigated in the laboratory under controlled humidities, and in the field under ambient conditions. For M. lemniscata, which has a relatively small clutch (8–30 eggs) and remains in the cocoon for approximately 9–10 months, removal of the cocoon had no effect on water loss from the egg stage, nor did it adversely affect hatching or molting success. Cocoon removal did, however, significantly affect water loss and, consequently, survival in the spiderling stage at all humidities in the laboratory and in the field. The importance of the cocoon for survival is probably related to the unusually long time M. lemniscata spiderlings spend in the cocoon overwintering. For A. aurantia, which has a substantially larger clutch size (300–1400 eggs) and remains in the cocoon for a shorter 6–7 months, cocoon removal had no effect on water loss, egg hatching success, molting success, nor spiderling survival. The lack of an effect suggests that other factors (e.g., relative humidity at the oviposition site, or a large clutch size) may be more important in controlling water loss for A. aurantia.     

Cotesia orobenae (Hymenoptera: Braconidae), a Gregarious Endoparasitoid of Evergestis rimosalis (Lepidoptera: Pyralidae), and Hyperparasitoids in Virginia Brassica Crops

Cotesia orobenae Forbes (Hymenoptera: Braconidae) is a gregarious endoparasitoid of the cross-striped cabbageworm (Evergestis rimosalis Guenée) (Lepidoptera: Pyralidae), a minor pest of Brassica crops in Virginia. Cocoons of C. orobenae were collected, weekly, from field plots of broccoli and cabbage crops in Whitethorne, Montgomery County (VA) during 1989 and 1990. Counts were also made of the eggs and larvae of E. rimosalis to determine the pest′s seasonality. In January 1991, C. orobenae cocoons were collected from the held plots to see if C. orobenae overwinters within cocoons. In May 1993, trap plants bearing E. rimosalis larvae were placed among the field plants weekly to determine if C. orobenae adults are present and ovipositing early in the season. Field counts of E. rimosalis eggs and larvae in 1990 indicated the occurrence of three generations off. rimosalis from June to October, but parasitism of larvae on trap plants by C. orobenae in May 1993 suggested that the parasitoid is present in mid-May. C. orobenae oviposits in first through third instar E. rimosalis and the resulting larvae feed internally on the developing host, exit the fourth (last) instar host, and spin cocoons in a loose mass. Mean brood size of 118 C. orobenae field-collected cocoon masses was 9.2 ± 4.4 cocoons per mass. Out of 1085 cocoons, adult C. orobenae emerged from 77%, hyperparasitoids emerged from 7%, and nothing emerged from 16%. Females accounted for 73% of the emerging C. orobenae adults. Data from field-collected cocoons suggest that C. orobenae enter diapause in late September and overwinter in cocoons. Six pupal hyperparasitoids were recovered: Isdromas lycaenae (Howard) (Ichneumonidae); Conura (=Spilochalcis) torvina (Cresson) and Conura (=Spilochalcis) albifrons (Walsh) (Chalcididae); Hypopteromalus tabacum (Fitch), Catolaccus cyanoideus Burks, and Trichomalopsis viridescens (Walsh) (Pteromalidae).     

Availability,uptake and regeneration of phosphate in mesocosms with varied levels of P deficiency

A population of cyclomorphic Bosmina coregoni was studied in Lake Östersjön, southwestern Sweden and results from field samples collected in 1988, 1989, 1990 and 1991 are presented. Animals collected in summer have remarkably higher carapace and prolonged antennule compared to what we call the normal morph. In 1991 the extreme morph reach its maximum body length, body height and antennule length in July to September. The occurrence of the extreme morph coincide with the hatching of the predaceous cladoceran Leptodora kindtii.The two morphs fluctuate in abundance and in relation to each other. In early spring only the normal morph occurred in the samples followed by a period of about two months when the two morphs were found together, in July only the extreme morph was found. In September the two morphs were again present in the lake. As has been shown for other cladoceran, the conspicuous carapace and antennule could be an adaptive response that decreases mortality due to invertebrate predation. Spectacular features like these are likely also accompanied by some sort of costs.     

Resting eggs, seasonal dynamics, and production of Bosmina longispina maritima (P. E. Muller) (Cladocera) in the northern Baltic proper

The seasonal dynamics and production of Bosmina longispina maritimawas studied both in the pelagic zone and bottom sediments ofthe northern Baltic proper in 1979. Resting eggs started tohatch in April–May, but the pelagic zone population didnot show signs of increase until August; this is typical forthe species in this area. Relatively few of those hatched inthe spring and early summer survived the low temperatures andlack of food below the thermocline. The population increasewas started with individuals hatched after the thermal stratificationweakened. From August until October the number of resting eggsin the sediment rose in direct response to their productionin the pelagic zone. A new hatching period began in October–November,but conditions in the pelagic zone were not suitable for theirfurther development. Such mistiming of hatching may, along withother factors, help to explain the large annual variations inthe productivity of B. longispina maritima in the Baltic. Sexualreproduction was found to be of relatively greater importancein the study area than in the areas where Bosmina reaches higherproductivity. This was assumed to be due to more intense selectionpressures in the prevailing conditions.     

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.     

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.     

Flow patterns around cocoons and pupae of black flies of the genusSimulium (Diptera : Simuliidae)

Flow patterns around structurally different cocoons and pupae of five species ofSimulium Latreille are described. Three features of the flow pattern common to all cocoons are; 1) a solenoidal vortex around the cocoon, 2) upward flow anterior (downstream) to the cocoon, and 3) one or two pairs of spiral-shaped vortices, which either touch or envelop the fill filaments of the pupa. The solenoidal vortex and the upward-spiralling, downstream vortices are common features of flow patterns around most bluff bodies submerged in a boundary layer. The proximity of the vortices to the fill filaments of all pupae suggests that these vortices are associated with gaseous exchange at the gill filaments.     

Intra- and interspecific predation on the cocoons of Erpobdella octoculata (L.) (Annelida: Hirudinea)

In laboratory experiments, eleven out of twenty-one invertebrate species and all of the three fish species tested fed on freshly laid flaccid cocoons of Erpobdella octoculata, though predation was only severe from a dytiscid beetle, E. octoculata and the fish species. There was a tendency for more cocoons to be eaten by starved than fed predators. Only snail species damaged one-week-old tougher cocoons.Only one out of 471 E. octoculata collected during their breeding season from the stony shores of two eutrophic lakes had a cocoon in its gut.The role of cannibalism of and interspecific predation on cocoons in the control and regulation of lake-dwelling populations of the erpobdellid is discussed, and assessed to be relatively unimportant. A more likely regulatory mechanism may involve high juvenile mortality from as yet unknown causes.     

Impact of cocoon predation and parasitism on endemic populations of the common pine sawfly, Diprion pini (L.) (Hymenoptera, Diprionidae) in different forest types

1 Predation and parasitism on litter‐buried cocoons of the common pine sawfly Diprion pini (L.) were compared in different forest types with endemic sawfly populations by field exposure of laboratory‐reared cocoons during three consecutive years (1993–1995). 2 The impact of cocoon predation was dependent on season and forest type. The highest predation (up to 95%) was found during autumn in forest stands with a dense understory vegetation. 3 Cocoon parasitism varied between year, season and forest type. The highest parasitoid attack was observed in pure pine forests with more or less barren soils, but did not exceed 24% of exposed cocoons. 4 Cocoons were exposed in small patches. Predators tended to exploit all cocoons of a patch, whereas parasitoids only attacked a few cocoons of a patch. Predation was similar on cocoons placed in the litter and those buried more deeply in the soil, whereas parasitism of soil‐buried cocoons was rare. 5 These results indicate that predators can have a remarkable potential for limiting endemic sawfly densities, if habitat conditions in a forest maintain their population and support their foraging behaviour. A notable effect of parasitoids on sawfly cocoons deposited in the litter is obviously restricted to typical pure and barren pine forests, but may play there a similar role as predation.     

Seasonal changes in age class structure and reproductive status of house mice on Marion Island (sub-Antarctic)

Feral house mice on sub-Antarctic Marion Island become reproductively active (males scrotal, females with perforate vaginas or pregnant) at an age >60 days and breed until death, which may occur at more than 13 months. Breeding is strongly seasonal; pregnant or lactating females were found only from October to May. A substantial proportion of mice old enough to breed in one summer overwinters to form a significant component of the breeding population the following summer but it is unlikely that any survive a second winter. The onset of breeding is closely synchronized with increasing day length but occurs about 2 months before mean temperature at the ground surface starts to increase significantly. Cessation of breeding is more closely associated with declining temperatures in late summer. For both males and females, the best correlation between reproductive activity and any of the temperature parameters measured was with average maximum temperature 1 cm above the ground. Competition for macroinvertebrate prey increases sharply in early winter due to high mouse numbers. The breeding season in 1991/1992 and 1992/1993 was at least 2 months longer than in 1979/1980, because the mice started breeding earlier, and stopped breeding later, in 1991/1993. The later cessation of breeding in 1991/1993 was despite the fact that there was a greater competition for macroinvertebrate prey, and that mean air temperatures during the early winter months were lower, than in 1979/1980.