Colonization, survival, and causes of mortality of Camerariu hamadryadella (Lepidoptera: Gracillariidae) on four species of host plants

Abstract.

• 1 The abundance, survival, and causes of mortality of Cameraria hamadryadella (Clemens) (Lepidoptera: Gracillariidae) were examined on four host plant species in Virginia, U.S.A. Quercus alba L. and Q.rubra L. are native within the geographic range of C.hamadryadella, and Q.robur L. and Q.benderi Baenitz are exotic. Q.robur is native to Europe, North Africa, and Asia and was probably introduced prior to 1850, and Q.benderi is of hybrid origin and introduced to cultivation before 1900. Q.alba and Q.robur are in the subgenus Lepidobalanus (white oaks), and Q.rubra and Q.benderi are in the subgenus Erythrobalanus (red oaks).
• 2 Larval mines of C. hamadryadella were abundant on both white oak species, including the exotic Q.robur, but were rare on host plants in the red oak subgenus. Un-hatched eggs of C.hamadryadella were not observed on red oaks. Other observations on host distribution indicate that C.hamadryadella is rarely found on red oaks. These observations are interpreted as circumstantial evidence that C. hamadryadella generally avoids ovipositing on red oaks.
• 3 Survival of C.hamadryadella to the adult stage was similar among all host species, but larvae tended to survive longer on hosts in the red oak subgenus. The observation of higher survival rates of early instar larvae on red oaks suggests that no nutritional or secondary chemical barrier reinforces the observed pattern of oviposition.
• 4 Significant differences in the distribution of the causes of mortality were detected between native and exotic host plant species. Larvae and pupae on native hosts were more likely to die because of predation, while those on exotic host plants were more likely to die because of parasitism and host feeding by adult female parasitoids. This pattern could arise because parasitoids prefer to forage on exotic host plants or because predators avoid foraging on exotic plants.
• 5 This study shows for C. hamadryadella that the only barriers to colonization and use of exotic hosts, in the white and red oak subgenera, are the presence of cues sufficient to stimulate oviposition and/or the absence of cues to deter oviposition. It also suggests that the presence of closely related native host plants in the region of introduction will increase the probability that exotic plants will be colonized by phytophagous insects.

     

Density-related foraging behaviour in Closterocerus tricinctus, a parasitoid of the leaf-mining moth, Cameraria hamadryadella

• 1 We examined the foraging behaviour of the parasitoid wasp, Closterocerus tricinctus (Ashmead) (Chalcidoidea: Eulophidae), as it visited larvae of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), in an outbreak population.
• 2 We tracked females of C.tricinctus, recording the time spent searching for mines and handling host larvae. The density of leaf-mines (host larvae) and their condition were recorded for each leaf visited. A subset of leaves visited by C.tricinctus was enclosed in fine mesh bags so that foraging success could be determined by rearing or dissection. The average density of mines and the average leaf-area mined was estimated for a random sample of leaves from each tree.
• 3 The selection of leaves upon which to forage appears to be density-dependent. C.tricinctus visits leaves with leaf-mine densities twice the average, and when switching leaves lands directly on leaf-mines 5 times more often than expected assuming random landings.
• 4 The total time spent foraging on a leaf, the average time spent handling hosts, and the total search time within leaves tend to decline on leaves with many hosts, but the observed declines are not statistically significant.
• 5 The proportion of leaf-mines visited within a leaf is strongly inversely density-dependent. 30% of visits to leaf-mines are re-visits and 29% of handling time is spent re-handling previously visited hosts. Furthermore, only 21% of visits to mines lead to successful parasitism. We suggest that self-interference and the avoidance behaviour of the host may reduce the number of visits of leaf-mines by C.tricinctus within a leaf.
• 6 The effect of the strongly inversely density-dependent foraging investment within leaves is to offset the observed density-dependent pattern of leaf visitation making the overall spatial pattern of visitation by C.tricinctus to mines of C.hamadryadella inversely density-dependent.
• 7 We suggest that the uncertainty of C.tricinctus surviving on multiply mined leaves because of density-dependent host mortality due to intraspecific competition in high-density host populations, the rarity of high-density host populations, and the rarity of multiply-mined leaves in low-density host populations combine to select against an aggregative response within leaves by C.tricinctus.

     

Effects of the light environment on oviposition preference and survival of a leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), on Quercus alba L.

Abstract.  1. Oviposition preference and the survival and mortality of larvae of Cameraria hamadryadella (Lepidoptera) on the foliage of Quercus alba L. (Fagaceae) in sun and shade were examined. To separate the effects of the environment of foliation from that of rearing, C. hamadryadella were allowed to oviposit on oak saplings that foliated either in the sun or shade. Subsequently, trees were placed into the sun or shade and larval survival was monitored.
2. Trees that foliated in the shade had leaves that were 30% longer than those that foliated in the sun. However, the density of leaf mines did not differ between trees that foliated in the sun or shade.
3. Larval survival was higher in the shade than in the sun, but no difference in survival due to the site of foliation was detected. Differences in larval survival between the sun and shade environments could not be attributed to differences in natural enemy mortality. However, 'death by other causes' was higher for larvae reared in the sun than in the shade. In sunny environments photochemical processes may reduce foliage quality, which results in lower survival of C. hamadryadella larvae.
4. Ovipositing C. hamadryadella do not appear to discriminate between foliage produced in the sun or shade. This lack of discrimination, coupled with the fact that ovipositing C. hamadryadella are active after dark, accounts for the absence of a difference in abundance between sun and shade.
5. The results presented here, combined with other published studies, argue that different patterns of insect herbivory and abundance in relation to the light level experienced by host-plant foliage might be expected for species that oviposit either in the night or in the day.     

Factors favouring the development and maintenance of outbreaks in an invasive leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae): a life table study

1 Life tables were constructed for seven generations of the invasive horse‐chestnut leaf miner Cameraria ohridella in Switzerland, to assess the factors allowing the moth to build and maintain outbreak densities and to identify ‘gaps’ among their mortality factors that could be targeted by new control methods. The fecundity of females was calculated and the mortality factors affecting all developmental stages determined. 2 Females contained 106.6 eggs. In captivity, the mean number of eggs laid per female varied between 34 and 82 eggs, with a maximum of 184 eggs. Egg mortality was 18–75% and was mostly due to unknown factors. 3 The main mortality factors affecting larvae and pupae in the leaves were predation by birds and arthropods in all generation and leaf senescence in the autumn generation. Parasitoids killed less than 5% of the population. 4 Nearly 90% of overwintering pupae died in dead leaves, the majority of them being killed by earthworms and other leaf decomposers. As a result, the overwintering generations were the only ones showing a net rate of increase less than one. Non‐overwintering generations had net rates of increase between four and 11, allowing populations to build up from spring to autumn. 5 The persistence of high outbreak densities in Europe is explained by a combination of at least three factors: (i) high fecundity; (ii) multivoltinism; (iii) the unusually low impact of traditional natural enemies of leaf miners and particularly parasitoids. 6 Yearly variations in population densities could be at least partly explained by differences in larval mortality due to leaf senescence and intraspecific competition in the last generation in autumn.     

Host race formation in the leaf-mining moth Acrocercops transecta (Lepidoptera: Gracillariidae)

Genetic differentiation in ecological traits plays an important role in the reproductive isolation of phytophagous insects. The present study aims to elucidate the genetic changes involved during the process of host shifts, by combining analyses for (1) host adaptations, (2) pre‐ and postmating isolation, and (3) phylogeney among populations, using a leaf‐mining moth, Acrocercops transecta. This species is associated with Juglans ailanthifolia and Lyonia ovalifolia. Transplantation of the larvae demonstrated that the Juglans‐associated population completely failed to survive on Lyonia, whereas the Lyonia‐associated population survived on Juglans as well as on Lyonia. Females of respective host‐associated populations oviposited on their natal host plant only. An mtDNA‐based phylogeny clearly separated the Lyonia‐associated population from the Juglandaceae‐associated population, and indicated that the Lyonia‐associated population once evolved from the Juglandaceae‐associated population. These results indicate that the processes of host shifting from juglandaceous species to Lyonia involved genetic changes both in larval ability to use host plants and in host preference of females. The derived Lyonia‐associated population has retained the potential to assimilate the ancestral host, Juglandaceae. Mating between the two host‐associated populations was successful for both directions of crossing, and there were no significant differences in egg hatchability between hybrids and control crosses. No adults emerged when the F₁ hybrid larvae were maintained on Lyonia; however, on Juglans the F₁ hybrid larvae grew to adulthood as well as in the control, suggesting a lack of genomic incompatibilities between the two host‐associated populations. In conclusion, the results showed that the two host‐associated populations are host races that are partially reproductively isolated, and that the differences in performance and preference function as strong barriers against gene flow between the host races. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93 , 135–145.     

Parasitism of Cameraria ohridella (Lepidoptera, Gracillariidae) in natural and artificial horse-chestnut stands in the Balkans

1 The horse‐chestnut leafminer, Cameraria ohridella Deschka & Dimic, was discovered during a mass outbreak on planted horse‐chestnut trees, Aesculus hippocastanum, near Lake Ohrid in Macedonia. However, several peculiarities in the leafminer's biology and ecology, such as its inefficient control through natural enemies, indicate that it may be of exotic origin. 2 In this study, the parasitism of C. ohridella was examined in natural and artificial horse‐chestnut stands in South‐eastern Europe. The size and composition of the parasitoid complex of natural stands was comparable with that known from artificial stands. No specialist parasitoid species were detected in any of the samples. The only parasitoid species found exclusively in natural stands during this investigation were the braconid Colastes braconius and the eulophid Chrysocharis phryne. Both are known to be polyphagous on many leafminers in Europe and C. braconius has been often recorded from C. ohridella in artificial stands in previous studies. Parasitism levels varied within the same range (3.6–21.0%) throughout the entire region, in both types of habitats. 3 These results suggest that the parasitoid complex of C. ohridella in natural and planted horse‐chestnut stands in Europe is typically that of an exotic, recently introduced leafminer. Consequently, C. ohridella is probably not indigenous in the Balkans, but has been introduced to the area of its discovery from another continent.     

Techniques for continuous rearing and assessing host preference of a multivoltine leaf-mining moth, Acrocercops transecta (Lepidoptera: Gracillariidae)

Techniques for rearing the leaf‐mining moth Acrocercops transecta successively over several generations are described. Based on continuous rearing, some life historical parameters in A. transecta were determined. Because of its short generation time, successive rearing makes the moth suitable for mating experiments and a model organism to elucidate the mechanism of host‐associated speciation.     

Impact of the leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae) and bleeding canker disease on horse‐chestnut: direct effects and interaction

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Bottom-up and top-down effects in a tritrophic system: the population dynamics of Plutella xylostella (L.)–Cotesia plutellae (Kurdjumov) on different host plants

Abstract.  1. The effects of host-plant resistance on the population dynamics of the Diamondback moth, Plutella xylostella L., and its solitary parasitoid, Cotesia plutellae (Kurdjumov), were studied in replicated time-series experiments.
2. Host-plant resistance did not affect the equilibrial abundance of the Diamondback moth, but it affected the dynamics of Diamondback moth populations.
3. The mean population size of Diamondback moth showed no significant difference between Brassica rapa (a susceptible host plant) and Brassica napus (a partially resistant host plant) either in the presence or absence of the parasitoid.
4. Time-series analysis suggests that the dynamics of Diamondback moth on B. rapa were underpinned by delayed density-dependent processes. In contrast, the dynamics of the moth on B. napus were influenced by a direct density-dependent process.
5. Although measures of short-term parasitism showed a significantly higher rate of parasitism by C. plutellae on Diamondback moth feeding on B. napus compared with B. rapa , this individual performance does not translate into differences in the population dynamics. Analysis shows no significant difference in the persistence time of the population-level interaction between the host and parasitoid on the two different host plants.     

Maintenance of the phenology of the winter moth (Lepidoptera: Geometridae)

Adult winter moths ( Operophtera brumata (L.)) are active in late autumn or early winter. The eggs overwinter in the canopy of trees and hatch simultaneously with the bursting of host tree buds. Many young larvae disperse on the wind on silk strands. Larvae are polyphagus and feed until late spring when they pupate in soil or leaf litter. The duration of the egg and pupal stages is genetically determined and varies with latitude. The egg stage is long in the north and short in the south, while the pupal stage is short in the north and long in the south.
The literature on the ecology and physiology of winter moth is reviewed. The factors maintaining the unusual phenology are discussed. It is concluded that the larval stage is early because mature leaves of many host trees are unsuitable as food, because parasitism against later larvae is more intense, and because summer temperatures may be injurious to larvae. The adult period is late in the year so that the final stages of pupal development occur in cool conditions and so that adults emerge after most insect predators have ceased activity. Throughout most of the range retarding the adult emergence period would cause activity to be impeded by severe winter weather; in the south this is not so and it is suggested that eggs must be on the trees for a minimum period to ensure synchronization of egg hatch with bud burst. The protracted adult emergence period may be an adaptation reducing predation by birds.     

Spatio-temporal analysis of the population dynamics of the oriental moth,Monema flavescens (Lepidoptera: Limacodidae)

The comparative analysis of life tables of the oriental moth, Monema flavescens, obtained in 6 patches for 8 generations in 4 years revealed the following:

1. The ratio of maximum to minimum of cocoon density for each patch ranged from 5.34 to 22.50, each value being more than 3.20, the ratio for the whole study area.
2. The density change from adult to cocoon in the next generation caused most of the spatial variation in density change per patch. The rate of adult-to-cocoon population change showed spatial density dependence in some generations but not in others. When the change rate lacked spatial density dependence, it was the key-factor for spatial variation in adult density for the following few generations till the change rate recovered spatial density dependence. This was because of flooding, which killed the spatial density dependence existing potentially in the adult-to-cocoon change rate and damaged the same patches during the few successive generations.
3. The rate of population change from overwintered generation adults (summer ones) to first generation cocoons was not only the key-factor for the rate of throughout-the-year change but temporally density dependent in each patch. Therefore, the density for the whole study area is considered to fluctuate within a range. However, the strong equilibrium seen in the cocoon density for the whole study area was due to the floods that happened to occur when the density was near and at its maximum, and it is considered that such a strong equilibrium does not always occur.
4. In the population change from summer adults to first generation cocoons, temporal density dependence was found in all the patches, but it was found only in one patch in the population change from autumn adults to second generation cocoons. This was because the spatial density dependence seen in the former corresponded to the absolute density of adults, while that in the latter corresponded to the relative density.

     

Multiple host shifts between distantly related plants, Juglandaceae and Ericaceae, in the leaf-mining moth Acrocercops leucophaea complex (Lepidoptera: Gracillariidae)

Insect herbivores such as gall formers and leaf miners are often highly specialized and adapted to their respective natal host plants. Due to the specialization and adaptation, it is presumed that host shifts readily occur among closely related plant species. Leaf-mining moths, the Acrocercops leucophaea complex, consist of three species, A. leucophaea, A. defigurata, and A. transecta. Larvae of all the species of the complex feed on Juglandaceae plants, but A. leucophaea and A. transecta are also associated with an Ericaceae plant, which is quite distantly related to Juglandaceae. Such a host utilization as in this species complex is very rare among phytophagous insects. In the present study, we estimate the history of host shifts by reconstructing the phylogeny of the A. leucophaea complex using molecular data (partial sequence of mitochondrial COI, 12S rDNA, and ND5). Parsimony and maximum likelihood analyses indicated that the common ancestor of the A. leucophaea complex used Juglandaceae only, and that the association with Ericaceae has evolved in A. leucophaea and A. transecta independently. Parametric bootstrap analysis also supported multiple origins of the association with Ericaceae in this complex. These results imply that there are ecological and biochemical factors that promote host shifting between Juglandaceae and Ericaceae despite the two families being not closely related.     

Host-specific variation in Ieaf miner population dynamics: effects on density, natural enemies and behaviour of Stilbosis quadricustatella (Lepidoptera: Cosmopterigidae)

Abstract. 1. We compared high and low density populations of a leaf miner ( Stilbosis quadricustatella (Cham.)) on two host oaks to ascertain mechanisms influencing abundance. High density miner populations occurred on sand live oak, Quercus geminata (Fagaceae), found in homogeneous stands at inland and coastal sites. Quercus nigra , water oak, a patchily distributed inland species, supported a low density leaf miner population.
2. Average foliar nitrogen of Q.geminata was significantly lower than that of Q.nigra , and lad mining period on Q.geminata was correspondingly longer than on Q.nigra .
3. The average leaf area of Q.nigra was significantly greater than that of Q.geminata .
4. Parasitism was significantly greater in Q.geminata miner populations and predation was significantly higher in Q.nigra populations. However, parasitism and predation rates were roughly reciprocal so that overall larval mortality did not differ significantly between hosts.
5. In a laboratory experiment, pupal overwintering survivorship did not differ significantly between moist and dry treatments of the sand and loam soil types that typify Qgeminata and Q.nipra habitats, respectively.
6. Within-leaf miner density on Q.geminata significantly affected larval survivorship, parasitism and predation. Leaf area and within-leaf miner density were positively correlated.
7. We detected no host-patch area or isolation effect on miner densities on coastal Qgeminata and there was no significant gradient of local coastal conditions affecting larval survivorship or natural enemies.
8. Coastal leaf miner densities were significantly higher on oak patch edges than in interiors.     

Mitochondrial and microsatellite DNA markers reveal a Balkan origin for the highly invasive horse-chestnut leaf miner Cameraria ohridella (Lepidoptera, Gracillariidae)

Biological invasions usually start with a small number of founder individuals. These founders are likely to represent a small fraction of the total genetic diversity found in the source population. Our study set out to trace genetically the geographical origin of the horse-chestnut leafminer, Cameraria ohridella , an invasive microlepidopteran whose area of origin is still unkown. Since its discovery in Macedonia 25 years ago, this insect has experienced an explosive westward range expansion, progressively colonizing all of Central and Western Europe. We used cytochrome oxidase I sequences (DNA barcode fragment) and a set of six polymorphic microsatellites to assess the genetic variability of C. ohridella populations, and to test the hypothesis that C. ohridella derives from the southern Balkans (Albania, Macedonia and Greece). Analysis of mtDNA of 486 individuals from 88 localities allowed us to identify 25 geographically structured haplotypes. In addition, 480 individuals from 16 populations from Europe and the southern Balkans were genotyped for 6 polymorphic microsatellite loci. High haplotype diversity and low measures of nucleotide diversities including a significantly negative Tajima's D indicate that C. ohridella has experienced rapid population expansion during its dispersal across Europe. Both mtDNA and microsatellites show a reduction in genetic diversity of C. ohridella populations sampled from artificial habitats (e.g. planted trees in public parks, gardens, along roads in urban or sub-urban areas) across Europe compared with C. ohridella sampled in natural stands of horse-chestnuts in the southern Balkans. These findings suggest that European populations of C. ohridella may indeed derive from the southern Balkans.     

Linking warming effects on phenology,demography, and range expansion in a migratory bird population

Phenological changes in response to climate change have been recorded in many taxa, but the population‐level consequences of these changes are largely unknown. If phenological change influences demography, it may underpin the changes in range size and distribution that have been associated with climate change in many species. Over the last century, Icelandic black‐tailed godwits (Limosa limosa islandica) have increased 10‐fold in numbers, and their breeding range has expanded throughout lowland Iceland, but the environmental and demographic drivers of this expansion remain unknown. Here, we explore the potential for climate‐driven shifts in phenology to influence demography and range expansion. In warmer springs, Icelandic black‐tailed godwits lay their clutches earlier, resulting in advances in hatching dates in those years. Early hatching is beneficial as population‐wide tracking of marked individuals shows that chick recruitment to the adult population is greater for early hatched individuals. Throughout the last century, this population has expanded into progressively colder breeding areas in which hatch dates are later, but temperatures have increased throughout Iceland since the 1960s. Using these established relationships between temperature, hatching dates and recruitment, we show that these warming trends have the potential to have fueled substantial increases in recruitment throughout Iceland, and thus to have contributed to local population growth and expansion across the breeding range. The demographic consequences of temperature‐mediated phenological changes, such as the advances in lay dates and increased recruitment associated with early hatching reported here, may therefore be key processes in driving population size and range changes in response to climate change.     

The fine-scale population dynamics of spruce budworm: survival of early instars related to forest condition

Abstract.  1. A lagged, density-dependent relationship between survival of early instars and host-tree condition is revealed during outbreaks of spruce budworm, Choristoneura fumiferana Clem. Persistent damage to hosts leads to deterioration of the stand.
2. Resource limitation affects survival during early-instar dispersal of spruce budworm. Impediments to distinguishing these events with estimates of survival were overcome with a simple model that describes the dispersal and survival processes. The model was used to analyse a recent 15-year population series from Black Sturgeon Lake and two historical datasets from Green River, in Canada.
3. Defoliation-induced damage to the trees resulted in increased losses of spring-emerging larvae that are dispersing in search of feeding sites. Losses were further exacerbated by biotic factors such as maternal fecundity, rates of infection by the pathogen, Nosema fumiferanae , and by weather-related effects on the foraging period.
4. Survival of early-stage budworm larvae in persistent outbreaks declined and the likelihood of other density-related factors such as rate of mortality from natural enemies increased. These results may reconcile outstanding differences in interpretation of the role of the forest resource in spruce budworm population dynamics and point to a common process linking the dynamics of other well-known budworm species.     

Population structure,demography, and dynamics of mountain baboons: An interim report

Counts of 61 baboon troops (Papio cynocephalus ursinus) at four localities in the Drakensberg mountains confirmed earlier reports of a small mean troop size. This troop size of 22.49 animals changed neither with latitude nor elevation. Data from two of the sites suggested that population density increases from south to north, while a working assumption of 2.5 animals/ km² allowed us to set the population size at 7,540 animals, living in 335 troops. Both the adult sex ratio of 2.07 females/male and the immature/ adult female ratio of 1.17 were unaffected by troop size. Repeated counts from nine known troops revealed that the population is at equilibrium. © 1995 Wiley-Liss, Inc.     

Mortality factors affecting the leaf-mining stages of Phyllonorycter (Lepidoptera: Gracillariidae) on oak and birch 1. Analysis of the mortality factors

Mortality of life stages of Phyllonoycter species in mined leaves of oak and birch was investigated in a Cheshire locality during 1974. Phyllonotycter species are mostly bivoltine, but P. cavella is univoltine. Parasitism and host-feeding (predation) by Hymenoptera caused most mortality. Parasitism was heaviest in the first generations, with ectoparasitism predominating, although endoparasitism caused inore mortality in the second generations. The incidence of host-feeding increased to leaf-fall when more mines contained host-fed remains than either parasites or healthy Phyllonoycter . Host-feeding occurred particularly in the first three (sap-feeding) larval instars; ectoparasitism affected mainly the two subsequent (tissue-feeding) larval stages and pupae; and endoparasitism occurred in all stages although affected hosts are killed only from the fourth larval instar onwards. Highest mortality was suffered by tissue feeders in the first generation but by sap feeders in the second. Total mortality was greatest in second generation mines and, in a given generation, survival curves for Phyllonotycter were similar on both tree species. The observed density of second generation mines relative to first in the sample was higher than predicted from first generation mortality levels, and this is discussed.