Progressive restriction of host plant exploitation along a climatic gradient: the willow psyllid Cacopsylla groenlandica in Greenland

1. Host plant selection by the endemic willow psyllid Cacopsylla groenlandica was studied at eight sites in three locations along a N–S climatic gradient in west Greenland.
2. C. groenlandica oviposited and developed on four willow species: Salix glauca, S. arctophila, S. uva-ursi and S. herbacea. Development took place on female catkins of all species and on leaf shoots of S. glauca .
3. At sites in southern Greenland, C. groenlandica fully utilized all four willow species. However, there was progressive reduction of host plant species and tissues exploited as latitude increased, such that towards the northern limit of its range the psyllid became highly specialized, feeding only on female catkins of S. glauca , despite the presence of alternative hosts. Where plant growth was tightly constrained within a restricted growing season, the demands of synchronizing with more than one species may have become too great, resulting in specialization on a single host.
4. The probable mechanisms producing specialization were between-species differences in host plant phenology, linked to variation in the growing season, set against the partially non-overlapping geographical distribution patterns of the host willows.
5. A small degree of phenological asynchrony between host plants can be advantageous for the insect. Under favourable conditions the reproductive season can be extended by using a series of host plants with staggered phenologies. Some 'early' hosts also create predator-free space. However, as asynchrony increases, the advantages disappear as the insect becomes unable to exploit the full range of host options.
6. The importance of host plant synchrony in the life histories of arctic insects is discussed.     

Variation in resource exploitation along an altitudinal gradient: the willow psyllids (Cacopsylla spp.) on Salix lapponum

Three congeneric species of Salix -fedding psyllid Cacospsylla spp. Occur along an altitudinal gradient (988 1222 m. a.s.l.) in southern Norway. The 27-km transect covered the local altitudinal range of the three species. Each species showed a different characteristic altitudinal distribution: C. propinqua occurred over the entire altitudinal range of the transect C. palmeni was restricted to higher altitude and C brunneipennis occurred manly at lower altitudes. This pattern of local altitudinal distribution mirrored the broader geographical distributions of these psyllids along latitudinal gradients. All three species developed on willow catkins, but also made use of leaves. Species differed in their use of leaves with >10% of sampled individuals of C. brunncipennis or C. Palmeni developing on leaves compared with up to 80% in C. propinqua. The proportion of C. propinqua nymphs developing on catkins increased with altitude and was also related to the temporal and spatial availability of catkins but was not a result of competitive displacement from catkins. Nitrogen concentration of host plants did not affect the proportions of C propinqua on leaves or catkins and was not related to adult size or psyllid density on catkins. Patterns or resource use are discussed in relation to differences in the distribution and abundance of the three species along the transect.     

Preference and performance are correlated in the spittlebug Aphrophora pectoralis on four species of willow

Abstract 1. There was a positive correlation between oviposition and feeding preferences and offspring performance in the spittlebug Aphrophora pectoralis Matsumura (Homoptera: Cercopoidea, Aphrophoridae) on four species of willow Salix sp. (Salicaceae) growing near Sapporo, Japan. Spittlebugs preferred rapidly growing shoots where performance was highest.
2. When the effects of shoot length were removed, egg densities on willow species were associated with offspring performance on three of four species. Egg densities and survival rates were low on Salix integra and Salix miyabeana . Survival rates were high on Salix sachalinensis , which had high egg densities, and Salix hultenii , which had low egg densities.
3. Aphrophora pectoralis formed mating aggregations almost exclusively on S. sachalinensis but then dispersed to other willow species to oviposit.
4. Nymphs dispersed from the oviposition site to feed on nearby shoots within the same plant but they did not disperse to other willow plants. Nymphs had the same preference for rapidly growing shoots as ovipositing females, so they were able to refine the maternal choice by moving to larger shoots near the shoot on which they had eclosed.
5. The spittlebugs were highly aggregated at all life stages so that even at high densities only a small proportion of the most vigorously growing shoots was utilised.     

柑橘木虱两地理种群的内共生菌群落组成及传菌能力

柑橘黄龙病（Huanglongbing, HLB）是经柑橘木虱Diaphorina citri传播的最主要柑橘病害之一, 危害严重时能对柑橘产业造成毁灭性的破坏。为了鉴定福建和海南2个地理种群柑橘木虱的内共生菌群落组成, 本研究对16S rRNA部分保守序列进行PCR扩增, 并利用特异性引物对不同内共生菌进行了感染率检测; 另外, 还通过人工接虫的方法, 探索柑橘木虱成虫在带黄龙病菌蕉柑Citrus reticulata cv. Tankan上的获菌能力, 以及带菌柑橘木虱成虫对黄岩蜜橘C. reticulata cv. Subcompressa的传菌能力。研究发现, 这2个地理种群的柑橘木虱含有相同的内共生菌组成, 包括α-Proteobacteria, Wolbachia spp., γ-Proteobacteria, mycetocyte symbionts, β-Proteobacteria, Oxalobacter和β-Proteobacteria, Herbaspirillum, 而且这2个地理种群柑橘木虱的4种内共生菌的携带率均在95%以上。柑橘木虱成虫在带菌蕉柑上饲菌28 d后, 带菌率可达到82%, 而带菌柑橘木虱成虫在黄岩蜜橘上传菌75 d后, 可导致橘树整体带菌。本研究为柑橘木虱的进一步研究和防虫治病途径提供了一些理论依据。     

Cospeciation of psyllids and their primary prokaryotic endosymbionts

Psyllids are plant sap-feeding insects that harbor prokaryotic endosymbionts in specialized cells within the body cavity. Four-kilobase DNA fragments containing 16S and 23S ribosomal DNA (rDNA) were amplified from the primary (P) endosymbiont of 32 species of psyllids representing three psyllid families and eight subfamilies. In addition, 0.54-kb fragments of the psyllid nuclear gene wingless were also amplified from 26 species. Phylogenetic trees derived from 16S-23S rDNA and from the host wingless gene are very similar, and tests of compatibility of the data sets show no significant conflict between host and endosymbiont phylogenies. This result is consistent with a single infection of a shared psyllid ancestor and subsequent cospeciation of the host and the endosymbiont. In addition, the phylogenies based on DNA sequences generally agreed with psyllid taxonomy based on morphology. The 3' end of the 16S rDNA of the P endosymbionts differs from that of other members of the domain Bacteria in the lack of a sequence complementary to the mRNA ribosome binding site. The rate of sequence change in the 16S-23S rDNA of the psyllid P endosymbiont was considerably higher than that of other bacteria, including other fast-evolving insect endosymbionts. The lineage consisting of the P endosymbionts of psyllids was given the designation Candidatus Carsonella (gen. nov.) with a single species, Candidatus Carsonella ruddii (sp. nov.).     

Cospeciation of Psyllids and Their Primary Prokaryotic Endosymbionts

Psyllids are plant sap-feeding insects that harbor prokaryotic endosymbionts in specialized cells within the body cavity. Four-kilobase DNA fragments containing 16S and 23S ribosomal DNA (rDNA) were amplified from the primary (P) endosymbiont of 32 species of psyllids representing three psyllid families and eight subfamilies. In addition, 0.54-kb fragments of the psyllid nuclear gene wingless were also amplified from 26 species. Phylogenetic trees derived from 16S-23S rDNA and from the host wingless gene are very similar, and tests of compatibility of the data sets show no significant conflict between host and endosymbiont phylogenies. This result is consistent with a single infection of a shared psyllid ancestor and subsequent cospeciation of the host and the endosymbiont. In addition, the phylogenies based on DNA sequences generally agreed with psyllid taxonomy based on morphology. The 3′ end of the 16S rDNA of the P endosymbionts differs from that of other members of the domain Bacteria in the lack of a sequence complementary to the mRNA ribosome binding site. The rate of sequence change in the 16S-23S rDNA of the psyllid P endosymbiont was considerably higher than that of other bacteria, including other fast-evolving insect endosymbionts. The lineage consisting of the P endosymbionts of psyllids was given the designation Candidatus Carsonella (gen. nov.) with a single species, Candidatus Carsonella ruddii (sp. nov.).     

The influence of the host plant on the population dynamics of Acizzia russellae (Homoptera: Psyllidae)

Abstract. 1. The role of the host plant, Acacia karroo , in the population dynamics of the indigenous non-pest psyllid, Acizzia russellae , was investigated.
2. Population levels of A.russellae in early summer were almost 10 times greater on the regenerative foliage of pruned trees (cut back in spring) than on normal trees. During late summer and winter, populations declined far more slowly on pruned trees, so that very much higher numbers of psyllids were maintained on these trees through to the next spring. This finding, and others, suggest that natural enemies and climate are not major determinants in the population dynamics of this insect under normal conditions in the field.
3. Laboratory measurements of several chemical and physical characteristics of the foliage of pruned and normal trees did not reveal any differences which would account for the observed effects.
4. Measurements of organic nitrogen concentrations in the leaves of normal A. karroo trees in the field showed significant correlations between psyllid numbers and leaf nitrogen, but only during summer in November and January.
5. Psyllid performance in the laboratory (duration of life cycle, egg production, and dry weight of emergent females) was significantly better for psyllids reared on plants containing high amino nitrogen.
6. It is suggested that the availability of quantities of suitably high quality nutrients in the leaves of pruned A. karroo trees explains the "epidemic" population levels achieved by the psyllid on pruned plants. It seems that characteristics of the host plant impose the major limitation on psyllid population growth, which may explain the permanent low endemic population levels of this insect in the field.     

Host plant growth characteristics as determinants of abundance and phenology in jumping plant-lice on downy willow

1. Salix lapponum host plants at an upper altitudinal site differed significantly in size, structural density, phenology, growth performance, and spatial isolation from those growing at a lower site. 2. Plant differences were paralleled by significant differences in psyllid population density and phenology parameters, with psyllid population density, percentage of catkins occupied, and phenological development relatively lower or retarded at the upper site. Population densities at the upper site, nevertheless, remained high. 3. Plant measurements were good predictors of insect density, often explaining up to 73% of the variance in abundance among plants at a given site. 4. Sets of four plant characters identified by best subsets regression were better predictors of psyllid density and development than single factors, although differences were often not great and the combinations of characters selected by multiple regression sometimes differed from the best single predictors. 5. Best single predictors of psyllid density on catkins were measurements of plant size, particularly height, length, and basal stem diameter. Shoot density and catkin phenology were occasionally important but plant isolation and prior growth performance were less important. 6. By contrast with density, age structure of the psyllid population was predicted best from plant phenological measurements, notably catkin phenology.     

Factors Influencing Host Plant Choice and Larval Performance in Bactericera cockerelli

Among the many topics of interest to ecologists studying associations between phytophagous insects and their host plants are the influence of natal host plant on future oviposition decisions and the mechanisms of generalist versus specialist host selection behavior. In this study, we examined the oviposition preferences, behavior and larval development of the tomato/potato psyllid, Bactericera cockerelli. By rearing psyllids with two distinct geographically-linked haplotypes on different host plants, we were able to examine the role of natal host plant and potential local adaptation on host plant usage. Choice bioassays among three host species demonstrated that psyllids from California had clear preferences that were influenced by natal plant. We further found that patterns in choice bioassays corresponded to observed feeding and movement responses. No-choice bioassays demonstrated that there is little to no association between development and host-plant choice for oviposition, while also indicating that host choice varies between haplotypes. These findings support the concept that mothers do not always choose oviposition sites optimally and also add support for the controversial Hopkins'' host selection principle.     

Phylogenetic characterization and molecular evolution of bacterial endosymbionts in psyllids (Hemiptera: Sternorrhyncha)

Most sternorrhynchan insects harbor endosymbiotic bacteria in specialized cells (bacteriocytes) near the gut which provide essential nutrients for hosts. In lineages investigated so far with molecular methods (aphids, mealybugs, whiteflies), endosymbionts apparently have arisen from independent infections of common host ancestors and co-speciated with their hosts. Some endosymbionts also exhibit putatively negative genetic effects from their symbiotic association. In this study, the identity of endosymbionts in one major sternorrhynchan lineage, psyllids (Psylloidea), was investigated to determine their position in eubacterial phylogeny and their relationship to other sternorrhynchan endosymbionts. Small-subunit ribosomal RNA genes (16S rDNA) from bacteria in three psyllid species (families Psyllidae and Triozidae) were sequenced and incorporated into an alignment including other insect endosymbionts and free-living bacteria. In phylogenetic analysis, all sequences were placed within the gamma subdivision of the Proteobacteria. Three sequences, one from each psyllid species, formed a highly supported monophyletic group whose branching order matched the host phylogeny, and also exhibited accelerated rates of evolution and mutational bias toward A and T nucleotides. These attributes, characteristic of primary (P) bacteriocyte-dwelling endosymbionts, suggested that these sequences were from the putative psyllid P endosymbiont. Two other sequences were placed within the gamma-3 subgroup of Proteobacteria and were hypothesized to be secondary endosymbionts. The analysis also suggested a sister relationship between P endosymbionts of psyllids and whiteflies. Thus, a continuous mutualistic association between bacteria and insects may have existed since the common ancestor of psyllids and whiteflies. Calculations using a universal substitution rate in bacteria corrected for endosymbiont rate acceleration support the idea that this common ancestor was also the ancestor of all Sternorrhyncha. Compared with other P endosymbiont lineages, the genetic consequences of intracellular life for some psyllid endosymbionts have been exaggerated, indicating possible differences in population structures of bacteria and/or hosts.     

Ovipositional preferences, damage thresholds, and detection of the tomato-potato psyllid Bactericera cockerelli (Homoptera: Psyllidae) on selected tomato accessions

The tomato-potato psyllid Bactericera [Paratrioza] cockerelli (Sulc) has recently caused losses exceeding 50% on fresh market tomatoes in California and Baja, Mexico by injecting a toxin that results in a condition known as 'psyllid yellows'. The objectives of this study were to: (i) document oviposition preferences on a range of tomato cultivars; (ii) determine threshold levels for psyllid densities that would cause psyllid yellows on tomatoes within the first three weeks following transplanting; and (iii) identify the most important 'psyllid yellows' symptoms that might be used in surveying and monitoring for this pest. Plant lines tested included the commonly-planted commercial cultivars 'Shady Lady' and 'QualiT 21', an older, previously commercial cultivar '7718 VFN', a common cultivar planted by consumers 'Yellow Pear', and a wild type plant accession, PI 134417. When given a choice, psyllids significantly preferred 'Yellow Pear' and avoided PI 134417 for oviposition. Under no-choice conditions psyllids laid significantly fewer eggs on PI 134417, but all the other plant lines were equally good substrates for laying eggs. Thus, oviposition preference is not likely to provide a functional management strategy in large plantings. On 'Shady Lady', psyllids preferred to oviposit on plants already infested with adults. On both 'Shady Lady' and '7718 VFN' oviposition was significantly greater on plants previously infested by nymphs as compared to uninfested control plants. This suggests that, at least for some cultivars, there is a physiological change in plant attractiveness following psyllid feeding. 'Yellow Pear' and 'QualiT 21' were relatively tolerant of psyllids, requiring 18 nymphs per plant to produce the disease symptoms. Only eight nymphs per plant were needed on 'Shady Lady' and '7718 VFN'. For all cultivars, the pest density showed strong correlations with measurements such as the number of yellowing leaves and leaflets and distorted leaves, which were as good as or better than the first factor extracted from principal component analysis. Therefore, such measurements have the potential to simplify field surveys.     

Can early loss of affiliates explain the coextinction paradox? An example from <Emphasis Type="Italic">Acacia</Emphasis>-inhabiting psyllids (Hemiptera: Psylloidea)

Recent models indicate that coextinction events should be numerous but empirical support for coextinction is paradoxically scarce. I investigated whether the coextinction paradox is due to affiliates becoming extinct well ahead of their hosts, with the disjunction masking the coextinction event. Using psyllids (Hemiptera: Psylloidea) on acacias (Mimosaceae: Acacia) in southern Sydney as an example system, I compared the density of affiliate species and the proportion of specialists on threatened versus secure hosts. I observed reduced psyllid species densities on the threatened hosts Acacia pubescens and A. prominens relative to common, sympatric acacias, consistent with the loss of affiliates ahead of host plant extinction. However, contrary to predictions, I found no support for reduced affiliate specialisation on the threatened hosts. Rather, proximity to closely related hosts may be a stronger determinant of the proportion of specialised psyllid species on a host. In addition, generalists, as well as specialists, may be lost from declining plants due to loss of familiarity with rarely-encountered host plants.     

Establishment of <Emphasis Type="Italic">Psyllaephagus parvus</Emphasis> and <Emphasis Type="Italic">P. perplexans</Emphasis> as serendipitous biological control agents of Eucalyptus psyllids in southern California

The free-living lemon gum psyllid, Cryptoneossa triangula Taylor, and the lerp-forming spotted gum psyllid, Eucalyptolyma maideni Froggatt (Hemiptera: Psyllidae) are invasive pests of eucalypts in California. In 2007, Psyllaephagus parvus Riek (Hymenoptera: Encyrtidae) was discovered parasitizing spotted gum psyllids and Psyllaephagus perplexans Cockerell was collected from lemon gum psyllids. While neither parasitoid species was purposely introduced, presence of the parasitoids was significantly associated with reduced intensity and duration of population peaks for both psyllid species. Spring peaks were reduced more than fall peaks. We estimated minimum rates of parasitism from the ratio of mummies to live nymphs. Higher parasitism was recorded in coastal than inland locations during the spring, while parasitism was similar for coastal and inland populations in the fall. Logistic regression models suggest parasitoids were the determining factor of psyllid population densities, although physical parameters, such as irrigation, may affect psyllid or parasitoid populations.     

Plant volatiles and density-dependent conspecific female odors are used by Asian citrus psyllid to evaluate host suitability on a spatial scale

We investigated how chemical cues derived from female Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae) and their host plants affect host acceptance choices by conspecifics. In four-choice cage and two-choice olfactometer assays, female psyllids avoided conspecific female cues in a density-dependent manner. However, odors from citrus plants actively damaged by psyllid feeding were attractive to conspecific females. When odors from feeding-damaged plants were presented simultaneously with odors from female D. citri, attraction of female conspecifics was no longer observed as compared with a clean air control in olfactometer assays. In subsequent experiments, D. citri females were released within arenas that contained actively feeding-damaged or non-damaged (control) citrus plants, each with previously psyllid-infested and uninfested young leaves. D. citri development is linked to the presence of these newly emerging leaves which is the only site of nymphal development. Female D. citri were initially attracted by the actively damaged plants as compared with non-damaged controls. After acceptance of plants that were actively damaged by feeding, D. citri females preferentially chose and settled on uninfested young leaves as compared with previously infested young leaves. A herbivore-induced plant volatile attractant and a female-specific odor repellent appear to be complementary foraging cues providing psyllids with information at two spatial scales: (1) the whole plant level for choosing a plant potentially harboring male conspecifics for mating, and (2) the within plant level to reduce intra-conspecific competition by identifying previously exploited resources.     

Experimental Infection of Plants with an Herbivore-Associated Bacterial Endosymbiont Influences Herbivore Host Selection Behavior

Although bacterial endosymbioses are common among phloeophagous herbivores, little is known regarding the effects of symbionts on herbivore host selection and population dynamics. We tested the hypothesis that plant selection and reproductive performance by a phloem-feeding herbivore (potato psyllid, Bactericera cockerelli) is mediated by infection of plants with a bacterial endosymbiont. We controlled for the effects of herbivory and endosymbiont infection by exposing potato plants (Solanum tuberosum) to psyllids infected with “Candidatus Liberibacter solanacearum” or to uninfected psyllids. We used these treatments as a basis to experimentally test plant volatile emissions, herbivore settling and oviposition preferences, and herbivore population growth. Three important findings emerged: (1) plant volatile profiles differed with respect to both herbivory and herbivory plus endosymbiont infection when compared to undamaged control plants; (2) herbivores initially settled on plants exposed to endosymbiont-infected psyllids but later defected and oviposited primarily on plants exposed only to uninfected psyllids; and (3) plant infection status had little effect on herbivore reproduction, though plant flowering was associated with a 39% reduction in herbivore density on average. Our experiments support the hypothesis that plant infection with endosymbionts alters plant volatile profiles, and infected plants initially recruited herbivores but later repelled them. Also, our findings suggest that the endosymbiont may not place negative selection pressure on its host herbivore in this system, but plant flowering phenology appears correlated with psyllid population performance.     

Can methane suppression during digestion of woody and leafy browse compensate for energy costs of detoxification of plant secondary compounds? A test with muskoxen fed willows and birch

Digestion and metabolism of woody and leafy browse requires detoxification of plant secondary compounds that can incur an energy cost. Browse, however, inhibits methane (CH(4)) production and therefore could offset some costs of detoxification. We measured an index of heat increment of feeding (HIFi) and CH(4) production in muskoxen (Ovibos moschatus) given a single test meal (at 10 g/kg BM(0.75)) composed of hay mixed with one of three browse species (Willow: Salix alaxensis, S. pulchra; Birch: Betula nana). Detoxification cost was estimated as HIFi of browse diet-HIFi of hay diet and CH(4) compensation as CH(4) production of hay diet-CH(4) production of browse diet. CH(4) compensation was noted in 47% of 15 trials in which a detoxification cost was evident; six trials were with woody browse and one with leafy browse. Separate controls were responsible for the difference in CH(4) compensation for leafy browse vs. woody browse. Detoxification costs for twigs and leaves of B. nana were underestimated because of their low digestibility. In only one of six treatments was CH(4) compensation documented for B. nana. We conclude that energy saved by CH(4) suppression was small (<6%) compared with detoxification costs.     

Relative importance of host tree species and environmental gradients for epiphytic species composition, exemplified by pyrenomycetes s. lat. (Ascomycota) on Salix in central north Scandinavia

According to the continuum concept of vegetation, variation in species composition is primarily determined by complex environmental gradients. Species-gradient relationships of ground-dwelling, independent organisms are studied at scales ranging from centimetres to continents. In this study we use a balanced data set for pyrenomycetes on Salix to address if how the current species-gradient paradigm needs to be modified to apply to assemblages of organisms that are dependent on other organisms for their existence. The data from a transect across central-north Scandinavia included variation along climatic gradients in oceanicity (from oceanic to continental vegetation sections), and temperatures (from south boreal to alpine vegetation zones) and among five common and widely distributed Salix host species ( Salix caprea agg., S. glauca ssp. glauca , S. lapponum , S. myrsinifolia agg. and S. pentandra ). Ten individuals of each Salix host species were selected and carefully examined for pyrenomycetes within each combination of section and zone. Data for 28 species in the 28 combinations of section, zone and host were subjected to ordination and constrained ordination analyses. Host species was the most important source of variation in species composition, followed by zone and section which are the same major regional gradients that are important to plants. We use examples to discuss the contribution of local ecological and substrate gradients to the high variation explained by host species, concluding that host specificity per se occurs for these partly parasitic fungi. We therefore suggest that in order to account for variation in composition of species assemblages with strong degree of host dependence, general rules for species-gradient relationships need to be extended by inclusion of host specificity as separate factor.     

Secondary Endosymbionts of Psyllids Have Been Acquired Multiple Times

Previous studies have established that psyllids (Hemiptera, Psylloidea) contain primary endosymbionts, designated as Carsonella ruddii, which cospeciate with the psyllid host. This association appears to be the consequence of a single infection of a psyllid ancestor with a bacterium. Some psyllids may have additional secondary (S-) endosymbionts. We have cloned and sequenced the 16S–23S ribosomal RNA genes of seven representative psyllid S-endosymbionts. Comparison of the S-endosymbiont phylogenetic trees with those of C. ruddii indicates a lack of congruence, a finding consistent with multiple infections of psyllids with different precursors of the S-endosymbionts and/or possible horizontal transmission. Additional comparisons indicate that the S-endosymbionts are related to members of the Enterobacteriaceae as well as to several other endosymbionts and insect-associated bacteria. Received: 2 May 2000 / Accepted: 8 June 2000