Host resistance in defoliated pine: effects of single and mass inoculations using bark beetle-associated blue-stain fungi

1 In 1996, 7000 ha of pine forests were defoliated by the pine looper Bupalus piniaria in south‐western Sweden. 2 The susceptibility of trees of different defoliation classes (0, 30, 60, 90 and 100% defoliation) to beetle‐vectored blue‐stain fungi was tested in inoculation experiments. Forty and 120‐year‐old Scots pine trees were inoculated with ‘single’, i.e. a few inoculations of Leptographium wingfieldii and Ophiostoma minus, two blue‐stain fungi associated with the pine shoot beetle Tomicus piniperda. The young trees were also ‘mass’ inoculated with L. wingfieldii at a density of 400 inoculation points per m² over a 60 cm stem belt. 3 Host tree symptoms indicated that only trees with 90–100% defoliation were susceptible to the mass inoculation. 4 Single inoculations did not result in any consistent differences in fungal performance between trees of different defoliation classes, regardless of inoculated species or tree age class. 5 Leptographium wingfieldii produced larger reaction zones than O. minus, and both species produced larger lesions in old than in young trees. 6 As beetle‐induced tree mortality in the study area occurred only in totally defoliated stands, mass inoculations seem to mimic beetle‐attacks fairly well, and thus seem to be a useful tool for assessing host resistance. 7 As even severely defoliated pine trees were quite resistant, host defence reactions in Scots pine seem to be less dependent on carbon allocation than predicted by carbon‐based defence hypotheses.     

Role of kin recognition in oviposition preference and cannibalism by the predatory mite <Emphasis Type="Italic">Gynaeseius liturivorus</Emphasis>

Animals often select oviposition sites to minimize the predation risk for eggs and juveniles, which are more vulnerable to predation than adults. When females produce eggs in clusters, the eggs and juveniles are likely to suffer from cannibalism. Although cannibalism among siblings is known to be lower than among non-siblings, there have been few investigations into the possibility that females select oviposition sites that reduce the risk of cannibalism for the offspring. To test this possibility, we examined oviposition preference by adult females of the predatory mite Gynaeseius liturivorus in response to the presence of her own eggs and to eggs of other females, offering plastic discs as oviposition substrates. Although females did not clearly show a preference for plastic discs on which they had oviposited, they avoided plastic discs on which other females had oviposited. When eggs of other females were artificially placed on clean plastic discs, adult female mites avoided these discs, suggesting that the eggs were used as cues for oviposition preference. Cannibalism among juvenile siblings was lower than among non-siblings. These observations show that adult females and juveniles of G. liturivorus discriminate kin relationships among conspecific individuals. Therefore, oviposition preference by adult female G. liturivorus may lead to the reduced risk of cannibalism among offspring.     

Putative source of the invasive Sirex noctilio fungal symbiont, Amylostereum areolatum, in the eastern United States and its association with native siricid woodwasps

Two genotypes of the fungal symbiont Amylostereum areolatum are associated with the invasive woodwasp Sirex noctilio first found in North America in 2004. S. noctilio is native to Europe but has been introduced to Australasia, South America and Africa where it has caused enormous losses in pine plantations. Based on nucleotide sequence data from the intergenic spacer region (IGS) of the nuclear ribosomal DNA, the A. areolatum genotypes found in North America are most similar to genotypes found in Europe, and not to genotypes from the southern hemisphere. Although two IGS strains of A. areolatum were found in North America it cannot be stated whether A. areolatum was introduced to North America from Europe once or twice based on our study. Genetic groupings formed by sequencing data were in most cases supported by vegetative compatibility groups (VCGs). Other siricid woodwasp species in the genus Sirex are native to North America. The North American native Sirex edwardsii emerging from the same tree as S. noctilio carried the same strain of A. areolatum as S. noctilio. The North American native Sirex sp. ‘nitidus’ collected outside the geographical range of S. noctilio carried a unique strain within A. areolatum. Our findings of A. areolatum in the native North American species, S. sp. ‘nitidus’, contrast with the previous view that A. areolatum was not present in North America before the accidental invasion of S. noctilio.     

Strong indirect interactions of Tarsonemus mites (Acarina: Tarsonemidae) and Dendroctonus frontalis (Coleoptera: Scolytidae)

Phoretic mites of bark beetles are classic examples of commensal ectosymbionts. However, many such mites appear to have mutualisms with fungi that could themselves interact with beetles. We tested for indirect effects of phoretic mites on Dendroctonus frontalis, which attacks and kills pine trees in North America. Tarsonemus mites are known to carry ascospores of Ophiostoma minus, which tends to outcompete the mutualistic fungi carried by D. frontalis. Experimental additions and removals of mites from beetles demonstrated that Tarsonemus propagate O. minus in beetle oviposition galleries. Furthermore, the abundance of Tarsonemus and O. minus tended to covary in nature. These results verified a strong mutualism between Tarsonemus and O. minus. Results also indicated that O. minus is an antagonist of D. frontalis: beetle larvae seldom survived in the presence of O. minus (compared to 83% survival elsewhere). Apparently, this is an indirect result of O. minus outcompeting the two species of mycangial fungi that are critical to beetle nutrition. Thus, Tarsonemus mites close a loop of species interactions that includes a commensalism (mites and beetles), a mutualism (mites and O. minus), asymmetric competition (O. minus and mycangial fungi), and another mutualism (mycangial fungi and beetles). This interaction system produces negative feedback that could contribute to the endogenous population dynamics of D. frontalis. Reproductive rate of Tarsonemus was more temperature‐sensitive than beetle generation time (which constrains the time for mite reproduction within a tree). This differential temperature sensitivity produces a narrow range of temperatures (centred at 27°C) in which mite reproduction per D. frontalis generation can attain its maximum of 100 mites/beetle. Consequently, seasonal oscillations in temperature are predicted to produce oscillations in the D. frontalis community, and climatic differences between regions could influence the community to dampen or exacerbate the cyclical outbreak dynamics of D. frontalis.     

Lack of fidelity revealed in an insect–fungal mutualism after invasion

Symbiont fidelity is an important mechanism in the evolution and stability of mutualisms. Strict fidelity has been assumed for the obligate mutualism between Sirex woodwasps and their mutualistic Amylostereum fungi. This assumption has been challenged in North America where the European woodwasp, Sirex noctilio, and its fungal mutualist, Amylostereum areolatum, have recently been introduced. We investigate the specificity of the mutualism between Sirex and Amylostereum species in Canada, where S. noctilio co-infests Pinus with native Sirex nigricornis and its mutualist, Amylostereum chailletii. Using phylogenetic and culture methods, we show that extensive, reciprocal exchange of fungal species and strains is occurring, with 75.3 per cent of S. nigricornis carrying A. areolatum and 3.5 per cent of S. noctilio carrying A. chailletii. These findings show that the apparent specificity of the mutualism between Sirex spp. and their associated Amylostereum spp. is not the result of specific biological mechanisms that maintain symbiont fidelity. Rather, partner switching may be common when shifting geographical distributions driven by ecological or anthropogenic forces bring host and mutualist pairs into sympatry. Such novel associations have potentially profound consequences for fitness and virulence. Symbiont sharing, if it occurs commonly, may represent an important but overlooked mechanism of community change linked to biological invasions.     

Field Evaluations of Subterranean Termite Preference for Sap-Stain Inoculated Wood

Few studies have focused on interactions between subterranean termites and the ophiostomatoid fungal associates of pine bark beetles or root feeding weevils. Field stake tests were employed at four locations throughout Mississippi to determine the feeding preference of subterranean termites for blue-stained, unstained, and partially decayed southern pine sapwood stakes. This study also utilized wood decayed by Gloeophyllum trabeum, a fungus previously shown to elicit a positive subterranean termite feeding response, as a positive control. Stakes inoculated with G. trabeum received significantly more attacks than all other treatments after 16 weeks. Of the stakes attacked by subterranean termites, stakes inoculated with Ophiostoma minus were degraded faster than any other treatment. Subterranean termite preference for stakes treated with either of two Leptographium spp. and the untreated negative controls did not differ; however, each was fed upon less than all other treatments. The feeding rate on stakes inoculated with O. ips and G. trabeum being fed upon by subterranean termites was not significantly different. These results represent the first evidence of wood containing non-structurally degrading fungi (O. ips and O. minus) eliciting a feeding preference from subterranean termites greater than that of decayed wood. The implications of these results are particularly relevant to pine forest ecology, nutrient cycling, subterranean termite control, and the utilization of blue-stained southern pine building products in the southeastern U.S.     

Reproductive strategy of a woodwasp with no fungal symbionts, Xeris spectrum (Hymenoptera: Siricidae)

Experiments were conducted to elucidate the reproductive strategy of the siricid woodwasp, Xeris spectrum, which carries no substantial symbiotic fungi in its body, in a comparison with the life cycles of two fungus-carrying siricid woodwasp species, Sirex nitobei and Urocerus japonicus, by considering ecological traits such as seasonal patterns of occurrence, spatial distribution of emergence on a tree, and oviposition activities. Part of the X. spectrum populations emerged in spring, during May and June, while others emerged in summer, during August and September, simultaneously with other siricid fungus-carrying woodwasps. The vertical distribution pattern of X. spectrum emergence holes on the trunk closely coincided with the emergence hole pattern of S. nitobei. X. spectrum laid few eggs on fresh logs, old logs, or on logs inoculated with potato dextrose agar, whereas on logs inoculated with Amylostereum chailletii or A. areolatum, X. spectrum females oviposited no less than 30%, on average, of their potential eggs. Moreover, the oviposition sites on these logs were concentrated near the Amylostereum inoculation positions. These results indicate that X. spectrum has evolved a life history that utilizes fungal symbionts of other woodwasp species without itself possessing any symbiotic fungus. Moreover, X. spectrum has evolved a dual reproductive system in that (1) some adults emerge in summer during the same emergence period as the fungus-carrying woodwasps and thereby oviposit on host trees already inoculated with fungi, and (2) other adults emerge the next spring and oviposit on trees that were inoculated with fungi a year earlier. Received: 16 January 1997 / Accepted: 14 July 1997     

Cytoplasmic incompatibility and multiple symbiont infection in the ash whitefly parasitoid,Encarsia inaron

Terrestrial arthropods are commonly infected with maternally inherited symbionts that cause reproductive incompatibilities between hosts with differing infection status. Such symbionts can have major effects on the efficacy of a biological control program if releases are comprised of mixtures of differentially infected individuals. In this study, the ash whitefly parasitoid, Encarsia inaron (Hymenoptera: Aphelinidae) from Arizona was surveyed for the presence of heritable bacterial symbionts; experiments were also performed to test for two phenotypes known to be caused by Encarsia symbionts—cytoplasmic incompatibility and changes in oviposition behavior and host use. E. inaron has successfully reduced ash whitefly to non-pest status in all three locations it has been released (California, Arizona, and North Carolina) and is also notable as one of the only Encarsia species that is not autoparasitic, with both male and female wasps developing as primary parasitoids of whiteflies. We show that E. inaron is infected with both Wolbachia and Cardinium. While there was no effect of the symbionts on oviposition behavior or host use, crosses between doubly infected male wasps and uninfected females resulted in a severe reduction in the number of female offspring; male offspring production was unaffected. This study thus serves as a further warning that ascribing a phenotype to a symbiont with confidence depends on eliminating the possibility of a mixed infection, and establishes E. inaron as a useful model for dissecting Wolbachia–Cardinium interactions.     

Ophiostomatoid fungi can emit the bark beetle pheromone verbenone and other semiochemicals in media amended with various pine chemicals and beetle-released compounds

Fungal volatile compounds can mediate fungal-insect interactions. Whether fungi can emit insect pheromones and how volatile chemicals change in response to chemicals the fungi naturally encounter is poorly understood. We analyzed volatiles emitted by Grosmannia clavigera (symbiont of the mountain pine beetle) and Ophiostoma ips (symbiont of the pine engraver beetle) growing in liquid media amended with compounds that the fungi naturally encounter: (−)-α-pinene, (+)-α-pinene, (−)-trans-verbenol, verbenone, or ipsdienol. Nine volatile compounds were identified among the fungal and amendment treatments. Volatiles qualitatively and quantitatively differed between fungal species and among amendment treatments. The bark beetle anti-aggregation pheromone (−)-verbenone was detected from both fungi growing in (−)-trans-verbenol-amended media. G. clavigera and O. ips can emit beetle pheromones and other beetle semiochemicals, suggesting that ophiostomatoid fungi could contribute to the chemical ecology of bark beetles. However, such contributions could be modulated by the presence of other environmental chemicals.     

Fidelity Among Sirex Woodwasps and Their Fungal Symbionts

We report that associations between mutualistic fungi and their economically and ecologically important woodwasp hosts are not always specific as was previously assumed. Woodwasps in the genus Sirex engage in obligate nutritional ectosymbioses with two species of Amylostereum, a homobasid\iomycete genus of white rot fungi. In the present study, the Amylostereum species and genotypes associated with three species of Sirex native to eastern North America and one relatively recent invasive Sirex from Europe were investigated by comparing intergenic spacer regions (IGS). Sirex spp. were sampled over 6 years from 23 sites in six US states, ranging from Maine in the northeast to Louisiana in the southeast, to obtain samples of Amylostereum from mycangia of adult females. Two of the native Sirex species (Sirex nigricornis and Sirex nitidus) were associated with either Amylostereum chailletii or Amylostereum areolatum, refuting the hypothesis of strict species-specific relationships. However, the invasive Sirex noctilio and the native Sirex cyaneus were each collected with only A. areolatum or A. chailletii, respectively, although S. noctilio was associated with two different IGS genotypes of A. areolatum and S. cyaneus occurs sympatrically with the other native Sirex. In Pinus, the preferred host tree of S. nigricornis and S. noctilio, these species co-occurred in 25.9 % of trees sampled, and horizontal transmission of fungal strains from S. noctilio to S. nigricornis was documented, although only in one tree. The extent that further spread and establishment of S. noctilio will alter the composition of symbionts carried by native Sirex is unknown but will depend in part on the degree of flexibility in these host–symbiont associations.     

Do container size and predator presence affect Culex (Diptera: Culicidae) oviposition preferences?

Organisms with complex life cycles typically do not exhibit parental care. Hence, the ability of adult females to choose quality oviposition sites is critical for offspring success. Gravid females of many insect taxa have the capability to detect environmental conditions in water-holding containers (e.g., resource level, presence of competitors or predators) and to choose the sites that are most suitable for offspring growth and development. Mosquitoes may also detect physical container characteristics related to water permanence such as surface area, volume, or container size, and some species such as those in the genus Culex have been shown to prefer larger containers. However, predators may also preferentially colonize larger containers; thus, ovipositing females may face decisions based on cues of site quality that balance the costs and benefits for offspring. We used a field experiment to evaluate the oviposition preferences of two Culex species in response to experimental container size and predator abundances within the containers. We found that both species avoided ovipositing in the largest containers, which have high abundances of Chaoborus sp. and dragonfly larvae (predators). However, the container size most commonly chosen for oviposition (15-L buckets) also had high mean abundance per liter of dragonfly larvae. These results suggest either prey naiveté or reduced vulnerability of these species to dragonflies compared to Chaoborus sp. Other potential mechanisms for the observed patterns are discussed.     

Oviposition site selection of the Alcon blue butterfly at the northern range margin

Female oviposition decisions in insects may strongly affect offspring growth and survival, and thus determine population performance. In this study, we examined oviposition site selection in the xerophilous ecotype of the endangered myrmecophilous butterfly Phengaris (=Maculinea) alcon (Lepidoptera, Lycaenidae) in Estonia, at the northern distribution margin of both the butterfly and its host plant, Gentiana cruciata L. Egg distribution on individual host plants appeared to be highly uneven: plants carrying high egg loads contrasted to a high proportion of host plants without any or with a few eggs. Host plant use for oviposition was strongly dependent on plant characteristics and environmental context. Host plant height relative to the surrounding vegetation rather than the absolute height of host plants was a key factor determining the use of particular host plant individuals for oviposition. In particular, plants protruding above surrounding vegetation had a higher probability of being used for oviposition, and carried more eggs. Additionally, the number of eggs laid on individual host plants was positively associated with the presence of flowers and the number of shoots. More aggregated host plants received fewer eggs than those with less conspecifics around. Feeding damage by wild herbivores, found in a substantial proportion of the butterfly’s host plants, strongly reduced the number of eggs on individual plants. Our results underline the need to assure that butterfly’s host plants do not become overgrown by surrounding vegetation. Best practices for opening vegetation around host plants may need further studies that explicitly account for butterfly’s host ants—their abundance in relation to vegetation height and their response to opening vegetation.     

Effect of associated fungi on the immunocompetence of red turpentine beetle larvae,Dendroctonus valens (Coleoptera: Curculionidae: Scolytinae)

Abstract Dendroctonus–fungus symbioses are often considered as the ideal model systems to study the development and maintenance of ectosymbioses, and diverse interactions, including antagonism, commensalism and mutualism, have been documented between these organisms. The red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae) is a pine‐killing invasive beetle in northern China. Fungi species Ophiostoma minus, Leptographium sinoprocerum, L. terebrantis and L. procerum were associated with this bark beetle. Antagonistic interactions between D. valens and its associated fungi, such as O. minus and L. sinoprocerum, have been demonstrated, but the underlying causes of this phenomenon are unknown. Here, we first found the two tested fungi species retarded the net weight gain of D. valens larvae after completing 3‐day feeding on their media. Furthermore, we provide direct evidence indicating the effect of associated fungi on the immunocompetence of D. valens larvae to explain the documented antagonism. Our results showed that the activity of phenoloxidase and total phenoloxidase in D. valens larvae were significantly upregulated by two strains of associated fungi, O. minus and L. sinoprocerum as compared with the controls. The phenoloxidase ratio increased significantly in the larvae which had fed for 3 days on media inoculated with O. minus. Because insect immune defenses are costly to be deployed, these results could be explored as one of the underlying mechanisms of the documented antagonism.     

Female-Specific Specialization of a Posterior End Region of the Midgut Symbiotic Organ in Plautia splendens and Allied Stinkbugs

Many stinkbugs (Insecta: Hemiptera: Heteroptera) are associated with bacterial symbionts in a posterior region of the midgut. In these stinkbugs, adult females excrete symbiont-containing materials from the anus for transmission of the beneficial symbionts to their offspring. For ensuring the vertical symbiont transmission, a variety of female-specific elaborate traits at the cellular, morphological, developmental, and behavioral levels have been reported from diverse stinkbugs of the families Plataspidae, Urostylididae, Parastrachiidae, etc. Meanwhile, such elaborate female-specific traits for vertical symbiont transmission have been poorly characterized for the largest and economically important stinkbug family Pentatomidae. Here, we investigated the midgut symbiotic system of a pentatomid stinkbug, Plautia splendens. A specific gammaproteobacterial symbiont was consistently present extracellularly in the cavity of numerous crypts arranged in four rows on the midgut fourth section. The symbiont was smeared on the egg surface upon oviposition by adult females, orally acquired by newborn nymphs, and thereby transmitted vertically to the next generation and important for growth and survival of the host insects. We found that, specifically in adult females, several rows of crypts at the posterior end region of the symbiotic midgut were morphologically differentiated and conspicuously enlarged, often discharging the symbiotic bacteria from the crypt cavity to the main tract of the symbiotic midgut. The female-specific enlarged end crypts were also found in other pentatomid stinkbugs Plautia stali and Carbula crassiventris. These results suggest that the enlarged end crypts represent a female-specific specialized morphological trait for vertical symbiont transmission commonly found among stinkbugs of the family Pentatomidae.     

Host selection by Ibalia leucospoides based on temporal variations of volatiles from the hosts’ fungal symbiont

Insect parasitoids locate hosts via reliable and predictable cues such as volatile emissions from hosts and/or host plants. For insects that depend on mutualistic organisms, such as many wood‐boring insects, symbiont‐derived semiochemicals may represent a source of such cues to be exploited by natural enemies. Ultimately, exploitation of these signals may increase fitness by optimizing foraging efficiency. Female parasitoids of Ibalia leucospoides use volatiles from the fungal symbiont Amylostereum areolatum of their host Sirex noctlio to find concealed host eggs and young larvae within the xylem. We hypothesize that the temporal pattern of fungal emissions may indicate not only the presence of host larvae but also be used as a cue that indicates host suitability and age. Such information would allow female parasitoids to discern more efficiently between hosts within ovipositor reach from those already buried too deep into the xylem and out of reach. In this context, we assessed the behaviour of I. leucospoides females to volatiles of A. areolatum in a ‘Y’‐tube olfactometer at regular intervals over 30 days. We concurrently examined the fungal volatiles by headspace sampling through solid‐phase microextraction (SPME) followed by gas chromatography mass spectrometry (GC‐MS). We observed that females were attracted to volatiles produced by two‐week‐old fungal cultures, a period that matches when older larvae are still within ovipositor reach. Four chemical compounds were detected: ethanol, acetone, acetaldehyde and the sesquiterpene 2,2,8‐trimethyltricyclo[6.2.2.01,6]dodec‐5‐ene, with each compounds’ relative abundance changing over time. Results are discussed in the context of parasitoids fitness. Future studies involving electrophysiology, different collection techniques and further behavioural assays will help in identifying the compounds that convey temporal information to female parasitoids and have the potential for being used in integrated pest management programmes.     

Detection and avoidance of an entomopathogenic fungus by a generalist insect predator

Abstract.  1. Increasing evidence suggests that insects can assess their environment based on cues related to mortality risks to themselves or their offspring. Limited knowledge is available on such abilities in relation to entomopathogenic fungi, which can cause significant mortality in insect populations. In laboratory bioassays, the ability of the generalist predator Anthocoris nemorum L. (Heteroptera: Anthocoridae) to detect the presence of its natural enemy, the fungal pathogen Beauveria bassiana (Balsamo) Vuillemin (Ascomycota: Hypocreales) was investigated.
2. Behavioural observations were conducted on adults of A. nemorum foraging in choice and non-choice arenas treated with conidia suspensions of B. bassiana or just the carrier (control). The arenas consisted either of nettle leaves or soil. Additionally, behaviours in response to sporulating nettle aphid cadavers compared with uninfected aphids or paper balls were evaluated on nettle leaves. An oviposition experiment was also conducted in choice arenas on conidia-treated and untreated nettle leaves.
3. Males and females detected and avoided contact with leaf surfaces inoculated with B. bassiana . Females that were forced to enter fungus-treated leaf surfaces were very reluctant to do so. When females encountered cadavers sporulating with B. bassiana they rapidly withdrew compared with harmless paper ball dummies. Soil inoculated with B. bassiana did not affect A. nemorum behaviour or residence time compared with control soil. Females inserted significantly more eggs in control leaf areas compared with areas treated with B. bassiana conidia.
4. All results suggest that A. nemorum detects and avoids the pathogen B. bassiana when it forages on host plants with which it is adapted but not on soil surfaces. The adaptive significance of detection of entomopathogenic fungi is discussed.     

The fungal matrices of <Emphasis Type="Italic">Ophiostoma ips</Emphasis> hinder movement of the biocontrol nematode agent, <Emphasis Type="Italic">Deladenus siricidicola</Emphasis>, disrupting management of the woodwasp, <Emphasis Type="Italic">Sirex noctilio</Emphasis>

Deladenus (=?Beddingia) siricidicola (Tylenchida: Neotylenchidae) is the most effective biocontrol agent used against the invasive wood wasp, Sirex noctilio (Fabricius) (Hymenoptera: Siricidae). The nematodes feed and reproduce on the wood-inhabiting fungus, Amylostereum areolatum (Chaillet ex Fr.) Boidin (Russulales: Amylostereaceae) and parasitise larvae of S. noctilio. In the nematode biocontrol program, the nematodes are inoculated into herbicide-weakened ‘trap trees’. Recent declines in nematode parasitism of S. noctilio in Australia have coincided with an increased incidence of an exotic bark beetle, Ips grandicollis (Eichhoff) (Coleoptera: Curculionidae), attacking trap trees and vectoring a wood-inhabiting fungus, Ophiostoma ips (Rumbold) Nannfelt (Ophiostomatales: Ophiostomataceae), which may inhibit migration of the nematode within the tree to the detriment of S. noctilio biocontrol. Several in vitro and in vivo experiments were conducted to investigate the effect of fungal interactions on the ability of D. siricidicola to locate and reproduce on A. areolatum. Deladenus siricidicola showed preference to A. areolatum in the presence and absence of O. ips, but the presence of O. ips negatively affected the choice response and the number of eggs laid by the nematodes. Deladenus siricidicola was unable to survive and reproduce on O. ips. Results give a clearer understanding of the choice response of D. siricidicola in I. grandicollis infested trees, explaining the disruptive impact of bark beetles on biocontrol of S. noctilio, an effect that could extend from Australia to other important pine growing countries.     

Symbionts mediate oviposition behaviour in invasive and native woodwasps

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Effects of inoculation with native arbuscular mycorrhizal fungi on clonal growth of Potentilla reptans and Fragaria moschata (Rosaceae)

Many clonal plants live in symbiosis with ubiquitous arbuscular mycorrhizal (AM) fungi, however, little is known about their interaction with respect to clonal reproduction and resource acquisition. The effects of arbuscular mycorrhiza on the growth and intraclonal integration between ramets of two stoloniferous species were studied experimentally in a nutritionally homogenous soil environment. Two species coexisting at the same field site, Potentilla reptans and Fragaria moschata, were selected as model plants for the study. Pairs of their ramets were grown in neighbouring pots with each ramet rooted separately. Four inoculation treatments were established: (1) both mother and daughter ramets remained non-inoculated, (2) both ramets were inoculated with a mixture of three native AM fungi from the site of plant origin, (3) only mother or (4) daughter ramet was inoculated. The stolons connecting the ramets were either left intact or were disrupted. Despite the consistent increase in phosphorus concentrations in inoculated plants, a negative growth response of both plant species to inoculation with AM fungi was observed and inoculated ramets produced fewer stolons and fewer offspring ramets and had lower total shoot dry weights as compared to non-inoculated ones. A difference in the extent of the negative mycorrhizal growth response was recorded between mother and daughter ramets of P. reptans, with daughter ramets being more susceptible. Due to AM effect on ramet performance, and thereby on the source-sink relationship, inoculation also significantly influenced biomass allocation within clonal fragments. Physiological integration between mother and daughter ramets was observed when their root systems were heterogeneous in terms of AM colonization. These results hence indicate the potential of mycorrhizal fungi to impact clonal growth traits of stoloniferous plant species, with possible consequences for their population dynamics.     

Use of odours by Cycloneda sanguinea to assess patch quality

Adult ladybirds are likely to encounter various species of prey when foraging for oviposition sites. Optimal oviposition theory predicts that females should lay eggs in those sites that are the most suitable for offspring development. Therefore, factors that directly affect offspring mortality, such as the presence of predators and food, are expected to play an important role in the assessment of patch profitability by ladybird predators. Using a Y‐tube olfactometer, we tested whether the predatory ladybird Cycloneda sanguinea L. (Coleoptera: Coccinellidae) can use volatile cues to assess patch profitability and avoid predator‐rich patches. We assessed the foraging behaviour of C. sanguinea in response to odours associated with tomato plants infested with a superior prey, Macrosiphum euphorbiae Thomas (Homoptera: Aphididae), and with an inferior prey, Tetranychus evansi Baker and Pritchard (Acari: Tetranychidae), in the presence or absence of the heterospecific predator Eriopis connexa Mulsant (Coleoptera: Coccinellidae). Females of C. sanguinea significantly preferred plants infested by M. euphorbiae to plants infested by T. evansi and avoided odours emanating from plants on which E. connexa females were present. Our results show that C. sanguinea use volatile cues to assess patch profitability and to avoid patches with heterospecific competitors or intraguild predators.