Population studies of the beech leaf mining weevil (Rhynchaenus fagi) in Ireland and Scotland

Abstract. 1. Population budgets for Rhynchaenus fagi, based for the most part on samples of beech leaves taken in June, were constructed for a 10-year period (Northern Ireland) and a 3-year period (Scottish borders).
2. There were years of sustained population growth and sustained decline, but not enough evidence to suggest a cycle. Similar population levels were found at both sites in all but one year.
3. Key factors were associated with the adult stage, although egg mortality may also make a significant contribution to population change.
4. Unknown sources of mortality embodied in the residual mortality were also suspected of acting in a density dependent manner, compensating for variation in parasitism.
5. Death rate of larvae within mines was related to May temperature in Northern Ireland, but not so clearly in Scotland.     

The occurrence of an adult reproductive diapause in the univoltine life cycle of the beech leaf mining weevil,Rhynchaenus fagi L.

Summary

Juvenile hormone (JH) and α-naphthyl acetate (α-NA) esterase activity was measured on a daily basis during embryogenesis of the house cricket, Acheta domesticus. In eggs dissected from the lateral oviducts and embryos through blastokinesis, there were elevated levels of nonspecific JH esterase activity. The JH esterase activity could not be resolved from the α-NA esterase activity by gel filtration chromatography and the metabolism of both substrates was inhibited equally by 0,0-diisopropyl phosphorofluoridate (DFP). From blastokinesis through egg hatch, the JH esterase activity was maintained at relatively low levels and was resolved from the α-NA esterase activity by gel filtration. The α-NA esterase activity was inhibited by DFP while the JH esterase activity was relatively unaffected. Low JH titers in eggs must be maintained through blastokinesis for normal development. Elevated JH esterase activity in eggs during this period appears to have a functional role in the metabolism of maternal JH in the egg.     

A simple life table study based on development and mortality in the beech leaf mining weevil Rhynchaenus fagi L.

The developmental stages of the beech weevil, Rhynchaenus fagi L. are proposed as suitable material on which to base a practical study on some aspects of population dynamics. The life history of the weevil is explained, illustrating the convenience of the immature stages for the study of development and mortality in a population. The factors normally causing mortality during each of the developmental stages are described. Results obtained from the collection of a random sample of infested leaves are included as an example of the simplicity with which data can be obtained to illustrate the concept of the life table. The limitations of such a study as well as the possibilities for changing emphasis are briefly discussed.     

Bud burst and success of the beech weevil, Rhynchaenus fagi: feeding and oviposition

Abstract. 1. Overwintered and starved weevils fed on beech within 5 min of contacting the plant in spring, and oviposition occurred in less than 48 h after feeding.
2. Size of unlaid eggs was significantly reduced when feeding weevils were subsequently starved from beech for 3 days.
3. Date of bud burst did not affect the amounts of feeding or oviposition by weevils.
4. Weevils fed and oviposited preferentially among trees, and the same trees were preferred in two successive years. This preference was independent of bud burst date.     

Semiochemical attractants for the beech leaf‐mining weevil,Orchestes fagi

The beech leaf‐mining weevil, Orchestes fagi L. (Curculionidae: Curculioninae: Rhamphini), a pest of European beech, Fagus sylvatica L. (Fagaceae), was recently discovered infesting American beech, Fagus grandifolia Ehrh., in Nova Scotia, Canada. Adult O. fagi feed on both young and mature leaves of beech as well as on other species (e.g., raspberry, Rubus spp.), but oviposition and larval feeding are restricted to beech. Females oviposit in young developing beech leaves at the time of bud burst. We characterized volatiles emitted from buds, leaves, and sapwood of American beech and examined their potential as attractants alone or when combined with other weevil pheromones for O. fagi. We predicted that adults would be attracted to volatiles emitted from beech leaves, especially those emitted from bursting beech buds. Gas chromatography/mass spectrometry (GC/MS) analyses of volatiles collected from buds at pre‐ and post‐budburst identified two diterpene hydrocarbons, 9‐geranyl‐p‐cymene ( 1 ) and 9‐geranyl‐α‐terpinene ( 2a ), that were emitted in large amounts at the time of bud burst. Compound 1 significantly increased mean catch of males and total O. fagi (but not females) on sticky traps compared with unbaited controls. Y‐tube bioassays confirmed attraction of male O. fagi to bursting beech buds and compound 1 . Attraction of male O. fagi to 1 , emitted in large quantities from American beech, is likely adaptive because both oviposition and mating of O. fagi coincide with budburst. Our data suggest that traps baited with 1 may be useful for monitoring the spread of O. fagi in North America.     

Life table parameters affecting the population development of the woolly beech aphid, Phyllaphis fagi

A detailed study of the basic biological parameters affecting the population development of the woolly beech aphid, Phyllaphis fagi L. (Homoptera: Drepanosiphidae), was performed under field and laboratory conditions in order to obtain information about factors responsible for outbreak situations in forest nurseries. In the nurseries, the aphid eggs have found to hatch before budburst. The newly hatched nymphs were highly active in searching for feeding places, which resulted in higher nymph mortality in the first generation than in the following generations. Ten aphid generations were recorded in the nursery during one growing season. In the laboratory, development time, nymph mortality, reproduction, and adult lifetimes were assessed for P. fagi at four constant temperatures, i.e., 12, 15, 20, and 25 °C through four successive generations. Both temperature and aphid generation were found to affect life table parameters. The first generation, however, was less susceptible to changing temperatures than the following generations. Phyllaphis fagi reproductive effort was more dependent on temperature than on aphid generation. No significant difference was found in reproductive effort between apterous and alate females. The potential for outbreak situations therefore exists throughout the summer. The formation of winged morphs was not restricted to a few generations but rather continued for several months, with activity peaking in all years in mid-June. This had implications for migration to new seedlings in the nurseries. The highest r_m value was obtained at 20 °C. Nymphs born at higher temperatures were sometimes born deformed and unable to survive.     

Didymella fagi sp. nov. and its anamorphAscochyta fagi, causing the yellow leaf spot disease ofFagus crenata andQuercus mongolica var.grosseserrata in Japan

AnAscochyta fungus was obtained during a survey of leaf spot diseases ofFagus crenata in the mountains of Aomori prefecture in 1995. The pathogenicity of the fungus toF. crenata was confirmed by inoculation. Its teleomorph was first found on artificially infected leaves after the leaves were placed in an incubator at 5°C with a 12-h photoperiod (approximately 500lx, daylight strip lamps) for 4 mo. The fungus was found to be the causal agent of yellow leaf spots ofF. crenata andQuercus mongolica var.grosseserrata in the field. Comparison with similar fungi so far described from Fagaceae indicated that the anamorph isAscochyta fagi and the teleomorph is an undescribed species ofDidymella. The nameDidymella fagi is introduced for the teleomorph. Correlation between the two morphs has been proved by cultural, morphological and pathological studies.     

Anatomy of the stridulation apparatus of the beech leaf‐mining weevil and characterization of,and behavioral responses to,stridulation sounds

We investigated auditory signals and morphology of the stridulatory apparatus of the European beech leaf‐mining weevil, Orchestes fagi L. (Coleoptera: Curculionidae), an invasive herbivore now established in Nova Scotia, Canada, to determine their potential for enhancing survey tools to monitor the spread of the species in Canada. We recorded and described sounds produced by adult O. fagi, analyzed the morphology of the stridulatory mechanism for intersexual differences and asymmetry, and examined behavioral responses elicited in conspecifics by playback of stridulation recordings. Adult O. fagi produced sounds under three conditions: male in distress, female in distress, and male in the presence of female. Female distress chirps lasted significantly longer than male distress chirps and male chirps in the presence of females, but peak frequencies and mean number of chirps per s did not differ significantly among the three groups. Morphology of the stridulation structures in male and female O. fagi was compared using scanning electron microscopy. Orchestes fagi have an elytro‐tergal file‐ and scraper‐type sound production apparatus, through which sound is produced upon anterior motion of the abdomen. Female O. fagi have a ‘pars stridens’ that is longer and has more ridges than males. Width and number of ridges per length of pars stridens did not differ between the sexes. Evidence of asymmetry was found in male pars stridens, with the right side being longer than the left. Playback of recorded sounds to adult weevils suggests female O. fagi were repelled by sounds produced by distressed males.     

Identification of potential natural enemies of the pea leaf weevil,Sitona lineatus L. in western Canada

Sitona lineatus L., the pea leaf weevil, is an invasive pest of Pisum sativum L. (field pea) that has recently become established in Alberta, Canada. Adults consume seedling foliage and larvae feed on Rhizobium root nodules thereby reducing nitrogen fixation; both life stages can reduce yield. Eggs and adults are vulnerable to predators and parasitoids. In 2009, a series of experiments was undertaken to identify potential indigenous natural enemies of S. lineatus in southern Alberta. In three test arenas, eggs were exposed to starved ground beetles in no‐choice tests for 48 h. Egg debris was observed when eggs were exposed to Bembidion quadrimaculatum L., Microlestes linearis (LeConte), Bembidion rupicola (Kirby), Bembidion timidum (LeConte), Poecilus scitulus LeConte, and staphylinid beetles. Of the two most abundant carabid species, the smaller B. quadrimaculatum consistently removed significantly more eggs (94.6%) than the larger Pterostichus melanarius (17.4%). Similar results were observed in Petri dish and egg card tests. No egg debris was observed in tests with Pt. melanarius. Presence of the larger beetle, Pt. melanarius, resulted in a lower rate of egg removal by the smaller B. quadrimaculatum relative to the rates observed for B. quadrimaculatum alone. Intraguild predation of B. quadrimaculatum by Pt. melanarius was observed in 47% of tests. These results suggest that B. quadrimaculatum is a potential predator of S. lineatus eggs in field pea agroecosystems and should be considered for use in conservation biological control and integrated pest management programs.     

The habitat and feeding preferences of Littorina obtusata (L.) and L. mariae sacchi et rastelli

The vertical zonation of the flat periwinkles Littorina obtusata (L.) and L. mariae Sacchi et Rastelli is different, but overlaps. At a sheltered location on the Isle of Cumbrae, Scotland, L. obtusata, although most abundant on Fucus vesiculosus L. and Ascophyllum nodosum (L.) Le Jol. on the midshore, ranged over the entire shore from the Pelvetia zone down to the upper limits of Laminaria. Littorina mariae exhibited a much more restricted distribution being associated predominantly with Fucus serratus L. on the lower shore. In laboratory trials with adult plants, flat periwinkles found fucoid algae both more attractive and more palatable than other seaweeds. Littorina obtusata also preferred the reproductive receptacles of fucoid algae to their vegetative tissue. Tiny germlings of a diversity of seaweeds were readily consumed by L. obtusata. In choice experiments germling Ulva lactuca L. was preferred to adult U. lactuca, whereas adult Fucus serratus was preferred to germling F. serratus. Contrasting foraging behaviours between different species of flat periwinkle are discussed in relation to differences in their radular dentition.     

Monoterpenes emitted by the large pine weevil, Hylobius abietis (L.) feeding on Scots pine, Pinus sylvestris L.

Abstract. Attraction of the large pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae) to pine logs was enhanced by the presence of weevils already on the logs. We investigated the volatiles emitted by insects feeding on host trees to determine the mechanism of the attraction. Chemical analyses of the collected volatiles showed large amounts of monoterpenes. There were no qualitative differences in the collections, between Scots pine alone and pine plus feeding weevils, nor between males and females, mated or virgin. Strong quantitative differences in monoterpenes existed that were correlated to the surface of cut bark. Electroantennograms (EAGs) were recorded on male and female antennae stimulated by increasing doses of collected volatiles. EAGs for pine alone and pine plus weevils at the same stimulus loading in (a + P)-pinene gave superimposable curves. EAGs of male and female volatiles were also identical. These results suggest an attraction mediated by host plant allelochemicals rather than by an aggregation pheromone.     

Tillage affects the activity‐density,absolute density,and feeding damage of the pea leaf weevil in spring pea

Conversion from conventional‐tillage (CT) to no‐tillage (NT) agriculture can affect pests and beneficial organisms in various ways. NT has been shown to reduce the relative abundance and feeding damage of pea leaf weevil (PLW), Sitona lineatus L. (Coleoptera: Curculionidae) in spring pea, especially during the early‐season colonization period in the Palouse region of northwest Idaho. Pitfall traps were used to quantify tillage effects on activity‐density of PLW in field experiments conducted during 2001 and 2002. As capture rate of pitfall traps for PLW might be influenced by effects of tillage treatment, two mark‐recapture studies were employed to compare trapping rates in NT and CT spring pea during 2003. Also in 2003, direct sampling was used to estimate PLW densities during the colonization period, and to assess PLW feeding damage on pea. PLW activity‐density was significantly lower in NT relative to CT during the early colonization period (May) of 2001 and 2002, and during the late colonization period (June) of 2002. Activity‐density was not different between treatments during the early emergence (July) or late emergence (August) periods in either year of the study. Trap capture rates did not differ between tillage systems in the mark‐recapture studies, suggesting that pitfall trapping provided unbiased estimates of PLW relative abundances. PLW absolute densities and feeding damage were significantly lower in NT than in CT. These results indicate that NT provides a pest suppression benefit in spring pea.     

The consequences of alternating diet on performance and food preferences of a specialist leaf beetle

The food quality of a given host plant tissue will influence the performance and may also affect the preference behavior of herbivorous animals. As nutrient contents and defense metabolite concentrations can vary significantly between different parts of a plant and change over time, herbivores are potentially confronted with diet differing in quality even when feeding on a single plant individual. Here we investigated to what extent feeding exclusively either on young or old, mature leaves of Brassica rapa or on a mixed diet of young and old leaves offered in alternating order affects the larval performance, food consumption, and the host preference behavior of adult mustard leaf beetles, Phaedon cochleariae. Analyzing different leaf quality traits, we found lower water contents, no changes in C:N ratio but more than threefold higher glucosinolate concentrations in young compared to old leaves. Individuals reared on mixed diet performed as well as animals reared on young leaves. Thus, compared to animals feeding exclusively on highly nutritious young leaves, diet-mixing individuals may balance the lower nutrient intake by a dilution of adverse secondary metabolites. Alternatively, they may integrate over the variation in their food, using a previously assimilated resource for growth at times of scarcity. Animals reared on old leaves grew less and had a prolonged larval developmental time, although they showed increased consumption indicating compensatory feeding. Additionally, we found that experience with a certain diet affected the preference behavior. Whereas individuals reared exclusively on young leaves preferred young over old leaves for feeding and oviposition, we did not find any preferences by animals reared exclusively on old leaves or by females reared on alternating diet. Thus, in contrast to positive feedbacks for animals reared on young leaves, an integrative growth of diet-mixing individuals potentially leads to a lack of feedback during development. Taken together, our results suggest that different diet regimes can lead to comparable performance of mustard leaf beetles but experienced feedbacks may differ and thus convey distinct diet preferences.     

The feeding ecology of Stephen's lory and nectar availability in its food plants

Stephen's lory, Vini stepheni , is one of four landbirds which are endemic to Henderson Island and until recently, its habits and ecology were unknown. I studied the diet and feeding ecology of Stephen's lory with particular reference to nectar variation in dominant food plants. Stephen's lory feeds on nectar, pollen and fruits from a variety of plants with lepidopteran larvae also forming a large component of the diet. The two plants which were dominant nectar sources for Stephen's lory produced flowers with very different nectar yields. Scaevola sericea flowers contained larger maximum volumes of more concentrated nectar than Timonius polygamus flowers. However, Scaevola flowers were a less predictable nectar resource, and grew at much lower densities, than Timonius flowers. Lories fed on the two flower types at different frequencies and it was hypothesized that the net rate of energy gained from feeding from a similar sized patch of the two plant species was comparable. Although nectar volumes of Timonius polygamus flowers decreased from the early mornings up until late afternoons, the lories did not show a corresponding daily pattern of foraging behaviour. Nectar yields also increased with the age of Timonius flowers. Stephen's lory has probably been the least affected of its genus by adverse changes to its habitat, and any information gained on the species may be useful when conservation measures of other Vini species are addressed.     

Oviposition and feeding preferences of a flower-feeding weevil,Coelocephalapion aculeatum, in relation to conspecific damage to its host-plant

The oviposition and feeding preferences ofCoelocephalapion aculeatum Fall (Coleoptera: Apionidae), a host specific florivore ofMimosa pigra L. (Mimosaceae), were studied in relation to conspecific damage to its hostplant. Adults ofC. aculeatum cease ovipositing in inflorescences when the egg load reaches a number consistent with the larval carrying capacity of the inflorescence. The basis for this oviposition deterrence was examined by offering inflorescences damaged by adult feeding alone, larval feeding alone and a combination of adult feeding and oviposition. Adults preferred to oviposit on inflorescences which are not damaged by either adult feeding, larval feeding, or oviposition. No evidence for the existence of an oviposition deterring pheromone (ODP) was found. I suggest that the ability of a single host inflorescence to support the development of many larvae causes selection for the use of these oviposition deterring cues which can convey more quantitative information about the level of previous infestation than can ODPs. Adults fed a similar amount on damaged compared to undamaged inflorescences. These results assisted in the design of host range testing trials and allows predictions to be made about the effectiveness of this insect as a biological control agent.     

Regeneration of Lycium barbarum L. plants from leaf tissue,callus culture and callus protoplasts

The possibility of plant regeneration from leaf tissue, callus and callus protoplasts of Lycium barbarum L. has been studied. Leaf segments were cultured on B5 medium (Gamborg et al. 1968) containing 1.5 mg/1 6-benzylaminopurine and 0.5 mg/1 -naphthaleneacetic acid. Regeneration of shoots was initiated after 30 days of cultivation. Callus was obtained from leaf and internode tissues on MS medium (Murashige and Skoog 1962) containing 0.4 mg/1 of 2,4dichlorophenoxyacetic acid. Subsequently, callus was successfully subcultured on the same medium with 1 mg/l of 2,4-dichlorophenoxyacetic acid and 0.2 mg/l -naphthaleneacetic acid. Organogenesis in callus culture was obtained in the course of 40 days after transferring to TM-4 (Shahin 1984). Protoplasts were isolated from callus tissue grown in vitro using an enzymatic method. Cell colonies, minicallus formation and organogenesis were obtained. Shoots were rooted on Murashige and Skoog medium containing 0..1 mg/l -naphthaleneacetic acid. Regenerated plants were transferred to soil and were grown to maturity. Regenerated plants carried normal morphological traits.Abbreviations BA 6-benzylaminopurine - 2,4-D 2,4-dichlorophenoxyacetic acid - NAA -naphthaleneacetic acid - Zea zeatin - GA₃ gibberellic acid - MS Murashige and Skoog medium - B5 Gamborg medium     

Effect of feeding frequency on food intake and growth of Arctic charr, Salvelinus alpinus L.

Social interactions and dominance hierarchy effects are important factors governing rates of growth of Arctic charr, Salvelinus alpinus L. The effects of hierarchy were increased as access to food became more restricted, i.e. feeding frequency was reduced, but these effects could not be attributed to direct competition for food since fish were fed to satiation at each feeding period. The results suggest that, whilst some fish on the restricted feeding regime were able to maintain good rates of growth, feeding by the majority of the fish was inhibited by the presence of larger individuals. Due to the importance of these hierarchy effects it was not possible to demonstrate physiological adaptations in fish allowed infrequent access to food.     

Roles of odor, taste, and toxicity in the food preferences of lambs: implications for mimicry in plants

In the traditional sense, food ingestion consists of prehending, masticating, swallowing, and digesting plant matter. It is also possible to ingest plants without eating them. Volatile compounds are inhaled directly into the lungs and transported from the lungs into the bloodstream. Volatiles in high concentrations could presumably produce toxicosis, without an herbivore ever ingesting a plant in the customary sense. Volatile compounds may be aposematic, serving to warn potential predators of toxins in plants. We conducted three experiments to explore the roles of odor, taste, and toxicity in the food preferences of lambs. The first experiment determined if brief exposure to a novel odor followed by lithium chloride (LiCl)‐induced toxicosis caused lambs to avoid a familiar food that contained the odor. Lambs that sniffed coconut‐flavored barley and then received LiCl subsequently ate less coconut‐flavored barley than lambs that did not receive LiCl. The second experiment determined if lambs were deterred from eating a familiar food by the odor of Astragalus bisulcatus. A. bisulcatus is a malodorous (to humans) sulfur‐containing herb considered unpalatable and toxic. Neither odor nor intraruminal infusions of A. bisulcatus deterred lambs from feeding. The third experiment also determined how the degree of familiarity with the odor of A. bisulcatus, along with toxicosis, influenced preference of lambs for food with or without the odor of A. bisulcatus. Lambs with 8 d exposure to the odor but not given LiCl ate similar amounts of food, with and without the odor of A. bisulcatus, whereas lambs given LiCl showed a mild aversion to food with the odor during testing. Lambs with 1 d exposure to the odor but no LiCl ate similar amounts of food, with and without the odor, whereas lambs given LiCl showed a strong but transient aversion to food with the odor. Collectively, these findings show that lambs responded strongly to novel odors, but their response was transient and depended on the postingestive consequences of toxins and nutrients associated with odor inhalation. Thus, we submit that odor alone, in the absence of toxicosis or nociception, is not a deterrent to herbivores that continually sample foods and adjust intake based on the postingestive effects of toxins and nutrients. It also is unlikely that non‐toxic plants can mimic the odors of toxic plants to avoid herbivory (Batesian mimicry), unless the odors are indistinguishable by herbivores, again because herbivores constantly sample foods.     

The feeding height preferences of two goat breeds fed Grewia occidentalis L. (Tiliaceae) in the Eastern Cape,South Africa

Nguni goats and Boer goats, which are farmed together in the savannas of the Eastern Cape Province, depend on the same feed resources. The feeding height intake rates and preferences of one of their most preferred browse species, Grewia occidentalis L. (Tiliaceae), were studied in order to indirectly determine resource partitioning between these goat breeds on the basis of feeding height. The two hypotheses tested were that taller Boer goats feed at higher levels than do Nguni goats and that both breeds have feeding height preferences that are related to differences in intake rates. Four individuals of each breed were fed branches attached to a height foraging board. The two breeds showed no particular height preference in either season. Bite rates varied with feeding height, while bite sizes and intake rates increased with feeding height. Boer goats had lower bite rates compared to Nguni goats, although both breeds achieved similar bite sizes and intake rates. Bites sizes and intake rates for both breeds were higher in summer than in winter, possibly because more browse biomass was available per branch in summer than in winter since G. occidentalis is deciduous. There was no evidence to support that feeding height preference was related to differences in intake rates. It is concluded that Boer goats may compete for forage resources during times of resource limitation. Thus, Nguni goats might be a more favourable breed in semi-arid savannas because of their smaller body-size and, therefore, absolute nutrient requirements as well as their better reproductive performance compared to Boer goats.