Cocoon predation on diprionid sawflies: the effect of forest fertility

Predation by small mammals is thought to be one of the main regulators of outbreaking sawfly species. It has been suggested that predation may be lower in poor and dryish forests, and this is the reason why outbreaks often begin from this type of environment. We studied experimentally how fertility of the forest site affects cocoon predation experienced by two sawfly species, the common pine sawfly Diprion pini (Linnaeus) and the European pine sawfly Neodiprion sertifer (Geoffroy). We applied a fertilization treatment to selected pine-dominated barren forest sites in Finland, and 2–4 years later monitored predation on the sawfly cocoons in fertilized and control areas. The results did not support the idea that forest fertility was related to cocoon predation. We also could not verify that small mammal abundance was related to fertility of the forest. The most obvious pattern we observed was that the two sawfly species differed dramatically in predation experienced. N. sertifer has its cocoon phase in mid-summer and experienced only moderate predation (37%) whereas D. pini, with its cocoon phase in autumn, suffered from very heavy predation (96%). Our observations suggest that if predation is important in controlling the population dynamics of the species, its impact depends more on the sawfly species and season than on the fertility of the forest site. Received: 1 March 1998 / Accepted: 25 May 1998     

Larval mortality of pine sawflies [Hymenoptera: Diprionidae] in relation to pollution level: A field experiment

Larval mortality ofNeodiprion sertifer (Geoffroy),Diprion pini (L.) andGilpinia pallida (Klug) were studied in field experiments around a factory complex in southwestern Finland. Larval colonies were transferred on the shoots of Scots pines growing at different distances from the emission source. Larval mortality was highest near the factories. InN. sertifer, larval mortality caused by the nuclear polyhedrosis virus was higher and cocoon mortality caused by parasitoids was lower near the pollutant source. The most abundant parasitoid species wereSynomelix scutulata (Hartig) andLamachus eques (Hartig). 16–67% of theN. sertifer, 0–5% of theD. pini and the 73–100% ofG. pallida cocoons contained parasitoids oviposited during the larval period of the sawflies.      

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.     

Physiology of cold hardiness in cocoons of five earthworm taxa (Lumbricidae: Oligochaeta)

Earthworm cocoons are mostly found in the uppermost soil layers and are therefore often exposed to low temperatures during winter. In the present study, cocoons of five taxa of earthworms were investigated for their tolerance to freezing, melting points of cocoon fluids and dehydration of cocoons when exposed to a frozen environment. Embryos of the taxa investigated were freeze intolerant. The melting points of fully hydrated cocoon fluids were high (above –0.3°C) and thermal hysteresis factors were absent. Exposure to a frozen environment caused the cocoons to dehydrate drastically and dehydrated cocoons showed significantly lower super-cooling points than fully hydrated cocoons, reducing the risk of freezing for dehydrated cocoons. It is proposed therefore that the cold-hardiness strategy of the earthworm cocoons is based on dehydration upon exposure to subzero temperatures in the frozen environment. Cocoons of three surface-dwelling taxa, Dendrobaena octaedra, Dendrodrilus rubidus tenuis and Dendrodrilus rubidus norvegicus had lower supercooling points and survived frost exposure better than cocoons of two deeper-dwelling taxa, Aporrectodea caliginosa and Allolobophora chlorotica. One of the investigated taxa, D. r. norvegicus, was collected from a cold alpine habitat. However, it was not more cold hardy than the closely related D. r. tenuis collected from a lowland temperate habitat. D. octaedra was the most cold hardy taxon, its cocoons being able to withstand –8°C for 3 months and –13.5°C for 2 weeks in frozen soil.Abbreviations dw dry weight - fw fresh weight - SCP supercooling point     

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.     

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.     

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.     

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.     

Prey availability and eclosion-help of callow workers in the formicine ant Camponotus mus

Summary In the ant Camponotus mus the eclosion of imagines from the cocoons was shown to be controlled by nurse workers, which broke the cocoons and assisted the callows to energe. Short-term colony prey deprivation modified the eclosion-help behavior resulting in a delay in cocoon opening times. During long-term prey deprivation, no new cocoons were spun; the starved larvae grew and pupated when prey was again available. These findings as a colony strategy adapted to the changing condition of prey availabitity in a temperate habitat are discussed.     

Forest defoliator outbreaks under climate change: effects on the frequency and severity of outbreaks of five pine insect pests

To identify general patterns in the effects of climate change on the outbreak dynamics of forest‐defoliating insect species, we examined a 212‐year record (1800–2011) of outbreaks of five pine‐defoliating species (Bupalus piniarius, Panolis flammea, Lymantria monacha, Dendrolimus pini, and Diprion pini) in Bavaria, Germany for the evidence of climate‐driven changes in the severity, cyclicity, and frequency of outbreaks. We also accounted for historical changes in forestry practices and examined effects of past insecticide use to suppress outbreaks. Analysis of relationships between severity or occurrence of outbreaks and detrended measures of temperature and precipitation revealed a mixture of positive and negative relationships between temperature and outbreak activity. Two moth species (P. flammea and Dendrolimus pini) exhibited lower outbreak activity following years or decades of unusually warm temperatures, whereas a sawfly (Diprion pini), for which voltinism is influenced by temperature, displayed increased outbreak occurrence in years of high summer temperatures. We detected only one apparent effect of precipitation, which showed Dendrolimus pini outbreaks tending to follow drought. Wavelet analysis of outbreak time series suggested climate change may be associated with collapse of L. monacha and Dendrolimus pini outbreak cycles (loss of cyclicity and discontinuation of outbreaks, respectively), but high‐frequency cycles for B. piniarius and P. flammea in the late 1900s. Regional outbreak severity was generally not related to past suppression efforts (area treated with insecticides). Recent shifts in forestry practices affecting tree species composition roughly coincided with high‐frequency outbreak cycles in B. piniarius and P. flammea but are unlikely to explain the detected relationships between climate and outbreak severity or collapses of outbreak cycles. Our results highlight both individualistic responses of different pine‐defoliating species to climate changes and some patterns that are consistent across defoliator species in this and other forest systems, including collapsing of population cycles.     

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, ?? , ??–??.     

Effects of host age and cocoon position on attack rate by the hyperparasitoid,Eurytoma sp. [Hym.: Eurytomidae], on cocoons of the parasitoid,Cotesia (=Apanteles) glomerata [Hym.: Braconidae]

Host age preference of the hyperparasitoid,Eurytoma sp., attacking the cocoons of the gregarious parasitoid wasp,Cotesia (=Apanteles) glomerata L., was investigated in the field and laboratory. Under laboratory conditions of 20°C and L16D8 photoperiod.Eurytoma sp. parasitized cocoons of all ages, laying 7 to 10 eggs per cluster during a 24 h period. Field-collected cocoons also indicated that the host was parasitized regardless of its developmental stage. However, the mortality ofEurytoma sp. laid in cocoons on the day before host emergence was as high as 60%. Furthermore, progeny sex ratio (proportion males) reached 0.708 in eggs laid in the oldest cocoon clusters, whereas that for younger cocoons was strongly female-biased. Together, these facts suggest that older hosts are less suitable forEurytoma sp. than are younger ones, even though there was no significant decreasing tendency in the number of parasitized cocoons per cluster. In addition, the effect of cocoon position within a cluster was apparent, outer cocoons being more easily parasitized than inner ones. TheEurytoma sp. female oviposited at random on the free surface ofC. glomerata cocoons.     

Facts and myths of antibacterial properties of silk

Silk cocoons provide protection to silkworm from biotic and abiotic hazards during the immobile pupal phase of the lifecycle of silkworms. Protection is particularly important for the wild silk cocoons reared in an open and harsh environment. To understand whether some of the cocoon components resist growth of microorganisms, in vitro studies were performed using gram negative bacteria Escherichia coli (E. coli) to investigate antibacterial properties of silk fiber, silk gum, and calcium oxalate crystals embedded inside some cocoons. The results show that the previously reported antibacterial properties of silk cocoons are actually due to residues of chemicals used to isolate/purify cocoon elements, and properly isolated silk fiber, gum, and embedded crystals free from such residues do not have inherent resistance to E. coli. This study removes the uncertainty created by previous studies over the presence of antibacterial properties of silk cocoons, particularly the silk gum and sericin. © 2013 Wiley Periodicals, Inc. Biopolymers 101: 237–245, 2014.     

A hierarchical Bayesian model to estimate the unobservable predation rate on sawfly cocoons by small mammals

Predation by small mammals has been reported as an important mortality factor for the cocoons of sawfly species. However, it is difficult to provide an accurate estimate of newly spun cocoons and subsequent predation rates by small mammals for several reasons. First, all larvae do not spin cocoons at the same time. Second, cocoons are exposed to small mammal predation immediately after being spun. Third, the cocoons of the current generation are indistinguishable from those of the previous generation. We developed a hierarchical Bayesian model to estimate these values from annual one‐time soil sampling datasets. To apply this model to an actual data set, field surveys were conducted in eight stands of larch plantations in central Hokkaido (Japan) from 2009 to 2012. Ten 0.04‐m² soil samples were annually collected from each site in mid‐October. The abundance of unopened cocoons (I), cocoons emptied by small‐mammal predation (M), and empty cocoons caused by something other than small‐mammal predation (H) were determined. The abundance of newly spun cocoons, the predation rate by small mammals before and after cocoon sampling, and the annual rate of empty cocoons that remained were estimated. A posterior predictive check yielded Bayesian P‐values of 0.54, 0.48, and 0.07 for I, M, and H, respectively. Estimated predation rates showed a significant positive correlation with the number of trap captures of small mammals. Estimates of the number of newly spun cocoons had a significant positive correlation with defoliation intensity. These results indicate that our model showed an acceptable fit, with reasonable estimates. Our model is expected to be widely applicable to all hymenopteran and lepidopteran insects that spin cocoons in soil.     

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.     

First use of pheromones to detect phenology patterns and density relationships of pine sawflies in German forests

1 Monitoring studies of pine sawflies with pheromone traps were performed for the first time in Germany. Pheromone traps baited with species‐specific pheromone substances were installed in pine forests at different locations in Bavaria, Brandenburg and Lower Saxony during two years. 2 It was possible to track the flight phenology of Diprion pini, Gilpinia pallida and Neodiprion sertifer reliably and to get information about the number of generations of these species in 1997 and 1998. 3 A clear relationship between trap catch and population density could not be detected, but qualitative changes in trap catch caused by different density levels were observed. 4 For D. pini, trap catches were different among endemic populations of different forest types. Furthermore, catches of males reflected the results from the regular cocoon collections by foresters during the previous winter. 5 For N. sertifer, trap catches in endemic populations were well separated from trap catches on sites with higher sawfly densities. However, no significant correlation between trap catch and sawfly density or defoliation level could be found. These results suggest that the efficacy of the pheromone traps probably varied with biological features (sex ratio, density level, immigration) of the particular population.     

Fungi inhabiting knotwood of Pinus sylvestris infected by Porodaedalea pini

Abundance and diversity of fungi in naturally formed knots of Pinus sylvestris affected by Porodaedalea pini were investigated. Samples were taken from trees that were (i) affected, with internal heartwood decay and no conks, (ii) affected, with internal heartwood decay and conks and (iii) controls. The Illumina sequencing technology was used for amplification of DNA, sequencing and analysis. In total, 566,279 raw sequences were obtained from six samples. Sequences included 74% of culturable and 8.4% of non‐culturable fungi and 17.6% of organisms with no reference sequences in NCBI. Abundance of organisms in knotwood, measured as number of OTUs, ranged from 36,272 (29,506 for fungi) to 178,535 (177,484 for fungi) and differed significantly between two trees in a stand and between stands. The highest and lowest average number of fungal OTUs occurred in infected trees with no conks and in trees with conks, respectively. Number of taxa ranged from 171 to 415 and often differed significantly between two trees in one stand and between stands. Greatest diversity occurred in control trees. The number of fungal taxa shared by two trees in one stand was 67–152 and that shared by two stands was 51–141. The majority of fungi were Ascomycota. Those most common in pines affected by P. pini were Coniochaeta hoffmannii and C. fodinicola (19.65%–59.92%). Infundichalara microchona, Leotiomycetes spp. and Rhinocladiella atrovirens were also present. Another common species, Lecanora conizaeoides, occurred most often in control trees (0.30%–8.82%). Porodaedalea pini was detected only sporadically. Non‐culturable fungi were most frequent in the control trees. The greater average abundance and smaller average diversity of fungi in knots of trees infected by P. pini suggest that the pathogen successfully competes with some fungal species and does not inhibit the growth of survivors. Some fungi detected may contribute to production of natural biocides.     

Suspending cocoons to evade ant predation in Meteorus pulchricornis, a braconid parasitoid of exposed-living lepidopteran larvae

We tested the hypothesis that cocoon suspension by a thread in hymenopteran parasitoids is a defense tactic against predators, by comparing predation against suspended and non-suspended cocoons of the braconid wasp Meteorus pulchricornis on a Quercus phillyraeoides hedge on which workers of the common small ant Crematogaster matsumurai were foraging. The lost proportion of non-suspended cocoons, which were artificially attached to leaves of Q. phillyraeoides , markedly decreased with cocoon age, indicating a critical phase of predation on young cocoons. No suspended cocoons at age 1–12 h at the beginning of exposure were lost within 12 h, whereas more than 75% of same-aged non-suspended cocoons were lost in the same period. Predation against such young cocoons would be a strong force driving the evolution of cocoon suspension in parasitoids of exposed-living host insects.     

Effects of relative humidity on cocoon formation and survival in the braconid wasp Cotesia glomerata

Abstract The effects of relative humidity (RH) on cocoon formation and survival in the braconid parasitoid wasp Cotesia glomerata (L.) (Hymenoptera: Braconidae) are investigated under various humidity conditions (50, 75, 90, 95 and 100% RH) at 20 °C and under an LD 16 : 8 h photoperiod. The mortality rate at the time of egression from hosts under 100% RH is significantly higher than for other RHs. Cocoon clusters formed at 100% RH spread significantly more than those formed at 50, 75, or 90% RH. Developmental periods differ significantly among RHs under which wasps developed. The mean period from the egression from hosts to adult emergence is 8.7 days when developed at 50–95% RHs, and 8.0 days at 100% RH. The emergence rates of C. glomerata that are maintained under the same humidity conditions after egression from hosts are not significantly different among RHs. However, emergence rates from cocoons that are transferred from 100% RH to 50 and 75% RH are < 70%, although the rates are > 90% in most cases. Some wasps do not emerge from cocoons: more than 60% die after adult eclosion at all RHs; the relative frequency of adult deaths is approximately 90% at 50% RH. Relative humidity influences the cluster and cocoon status strongly: both good clusters and cocoons are formed at low RHs. Emergence rates from cocoons of different ranks are significantly different: the rates of low‐rank cocoons are low at low RHs. The survival of C. glomerata is affected strongly by RH through cocoon formation.