Species delimitation in downy mildews: the case of Hyaloperonospora in the light of nuclear ribosomal ITS and LSU sequences

Species definitions for plant pathogens have considerable practical impact for measures such as plant protection or biological control, and are also important for comparative studies involving model organisms. However, in many groups, the delimitation of species is a notoriously difficult taxonomic problem. This is particularly evident in the obligate biotrophic downy mildew genera (Peronosporaceae, Peronosporales, Oomycetes), which display a considerable diversity with respect to genetic distances and host plants, but are, for the most part, morphologically rather uniform. The recently established genus Hyaloperonospora is of particular biological interest because it shows an impressive radiation on virtually a single host family, Brassicaceae, and it contains the downy mildew parasite, Arabidopsis thaliana, of importance as a model organism. Based on the most comprehensive molecular sampling of specimens from a downy mildew genus to date, including various collections from different host species and geographic locations, we investigate the phylogenetic relationships of Hyaloperonospora by molecular analysis of the nuclear ribosomal ITS and LSU sequences. Phylogenetic trees were inferred with ML and MP from the combined dataset; partitioned Bremer support (PBrS) was used to assess potential conflict between data partitions. As in other downy mildew groups, the molecular data clearly corroborate earlier results that supported the use of narrow species delimitations and host ranges as taxonomic markers. With few exceptions, suggested species boundaries are supported without conflict between different data partitions. The results indicate that a combination of molecular and host features is a reliable means to discriminate downy mildew species for which morphological differences are unknown.     

Taxonomy of Mechanitis (F.) (Lepidoptera: Nymphalidae) from the West Colombian Andes: an Integrative Approach

Species identification in the butterfly genus Mechanitis (F.) (Lepidoptera: Nymphalidae) becomes difficult when it is based only on wing color patterns, a common practice in butterfly taxonomy. Difficulties in Mechanitis taxonomy are related to the widespread mimicry and polymorphism among species belonging to this genus. Species recognition and inventories of Mechanitis genus in geographic areas as the Andean region of Colombia are of particular interest and the use of more than one character for taxonomic identification is desirable. In this study, we included morphological, ecological, and mitochondrial DNA data to identify the occurring species in this region. Species of Mechanitis were studied from ecological, morphological, and molecular perspectives considering host plant identification, oviposition behavior, and life cycles under laboratory conditions. Immature morphology, patterns of wing color, and genital structures of adults were also studied. The genetic barcoding region of the cytochrome oxidase I mitochondrial gene was sequenced and used to verify the limits between species previously defined by the other characters and to validate its usefulness for species delimitation in this particular genus. The integrative approach combining independent datasets successfully allowed species identification as compared to the approach based on a single dataset. Three well-differentiated species were found in the studied region, Mechanitis menapis (Hewitson), Mechanitis polymnia (Linnaeus), and Mechanitis lysimnia (Fabricius). New valuable characters that could improve taxonomic identification in this genus are considered.     

First record of an apparently rare fig wasp feeding strategy: obligate seed predation

1. Fig trees require host‐specific agaonid fig wasps for pollination, but their figs also support numerous non‐pollinating fig wasps (NPFW) that gall fig tissues or develop as parasitoids. 2. Ficus microcarpa L. is widely naturalised outside its native range and the most invasive fig tree species. Seed predators are widely used for the biological control of invasive plants, but no obligate seed predatory (as opposed to ovule or fig wall galling) NPFW have been recorded previously from any fig trees. 3. Philotrypesis NPFW are usually regarded as parasitoids or ‘inquilines’ (parasitoids that also eat plant material) of pollinator fig wasps, but the present study provides evidence that Philotrypesis taiwanensis, a NPFW associated with F. microcarpa, is an obligate seed predator: (i) adults emerge from seeds of typical appearance, with a surrounding elaiosome; (ii) it shows no preference for figs occupied by fig wasp species, other than the pollinator; (iii) it only develops in figs that contain pollinated ovules, avoiding figs occupied by an agaonid that fails to pollinate; (iv) larvae are distributed in layers where seeds are concentrated and (v) it has a negative impact on seed but not pollinator offspring numbers. 4. Philotrypesis is a large genus, and further species are likely to be seed predators.     

Plant damage to vegetable crops by zoophytophagous mirid predators

The use of plant-feeding predators for biological pest control has traditionally been neglected, mainly due to the risk of them feeding on crop plants and causing economically significant damage. Yet, these predators offer advantages for biological pest control. They are mostly generalist predators that have an impact on several crop pests. They may also be able to establish on crops early in the growing season, when pests colonize them, and can remain on the target crop when prey is scarce. Therefore, management programs must seek to minimize risks while maximizing benefits. In vegetable crops, most of the literature on zoophytophagous predators has focused on four species: Dicyphus tamaninii, Dicyphus hesperus, Macrolophus pygmaeus and Nesidiocoris tenuis (Heteroptera, Miridae). The capacity of these species to produce crop damage in tomatoes varies. This damage has been related to relative predator-to-prey abundance, with damage increasing at high predator abundances and low prey densities. In this review, we analyze the use of these species in biological control programs and the associated benefits and risks. The differences in the damage caused by the four predatory species examined could not be attributed to either stylet morphology or saliva composition. However, feeding on specific plant structures where they may find the resources required for their development is what probably determines feeding damage. Understanding when and why these predators increase their feeding on plants or on certain plant parts is of crucial importance for integrating them in biological control programs.     

Plant feeding by a predatory mite inhabiting cassava

Plant feeding by arthropod predators may strongly affect the dynamics of bi- and tri-trophic interactions. We tested whether a predatory mite, Typhlodromalus aripo, feeds upon its host plant, cassava. This predator species is an effective biological control agent of Monoychellus tanajoa (the cassava green mite or CGM) a herbivorous mite specific to cassava. We developed a technique to detect plant feeding, based on the use of a systemic insecticide. We found that T. aripo feeds upon plant-borne material, while other predatory mite species, Neoseiulus idaeus and Phytoseiulus persimilis, do not. Subsequently, we measured survival of juveniles and adult females of T. aripo and N. idaeus, both cassava-inhabiting predator species, on cassava leaf discs. Survival of T. aripo was higher than that of N. idaeus. Thus, T. aripo was able to withstand longer periods of prey scarcity. Because CGM populations fluctuate yearly and are heterogeneously distributed within plants, plant feeding may facilitate the persistence of populations of T. aripo in cassava fields and its control of CGM outbreaks. This revised version was published online in July 2006 with corrections to the Cover Date.     

THE EVOLUTION OF HOST-PLANT USE AND SEQUESTRATION IN THE LEAF BEETLE GENUS PHRATORA (COLEOPTERA: CHRYSOMELIDAE)

Leaf beetles in the genus Phratora differ in host plant use and in the chemical composition of their larval defensive secretion. Most species specialize on either poplars or willows (family Salicaceae), but two species feed on birch (family Betulaceae). Phratora vitellinae utilizes salicylates from the host plant to produce its larval secretion, which contains salicylaldehyde, while other Phratora species produce an autogenous secretion. To reconstruct the evolutionary history of host plant use and the larval secretion chemistry in this genus, we sequenced 1383 base pairs of the mt cytochrome oxidase I gene for six European and one North American Phratora species and three outgroup taxa. Bootstrap values of the complete nucleotide sequence were 99-100% for six of eight nodes in the maximum parsimony tree. They were 71% and 77% for the two other nodes. The maximum parsimony tree and the maximum likelihood tree based on nucleotide sequence showed the same relationships as a maximum parsimony tree based on the amino acid sequence. Beetle phylogeny overlapped broadly with host plant taxonomy and chemistry, and it revealed historical constraints influencing host plant use. However, there was one host shift from the willow family (Salicaceae) to the birch family (Betulaceae). The use of host plant phenol glycosides for the larval defensive secretion evolved along the lineage that led to P. vitellinae. Phratora vitellinae feeds on the taxonomically widest range of host plants, which are characterized by moderate to high levels of salicylates. The results support the hypothesis that the use of salicylates for the larval secretion evolved twice independently in chrysomeline leaf beetles.     

Is Neoseiulus wearnei the Neoseiulus californicus of Australia?

Species of the family Phytoseiidae are known as predatory mites, some of them being used in crops to control mite pests, all around the world. Neoseiulus (=Cydnodromus) californicus is among the most commonly used Phytoseiidae species in biological control programs, especially in vineyards, orchards and vegetable fields. This species is distributed world-wide but has never been reported from Australia. On the other hand, specimens morphologically close to N. californicus have been assigned to a species called Neoseiulus wearnei, only reported from Australia. Investigations based on morphological and molecular comparisons were carried out to investigate whether these two taxa are conspecific. Morphological analyses showed no significant difference between specimens identified as N. wearnei and N. californicus. Similarly, genetic distances between these taxa were null, showing that all these specimens belong to the same species. Although it is not yet possible to conclude that all the specimens identified as N. wearnei are N. californicus, we can conclude that N. californicus is present in Australia. The information about the biology of N. californicus can thus now be applied to the Australian population of this species for biological control purposes.     

Implications of the changing phylogenetic relationships of Acacia s.l. on the biological control of Vachellia nilotica ssp. indica in Australia

Plant relationships have implications for many fields including weed biological control. The use of DNA sequencing and new tree building algorithms since the late 1980s and early 1990s have revolutionised plant classification and has resulted in many changes to previously accepted taxonomic relationships. It is critical that biological control researchers stay abreast of changes to plant phylogenies. One of the largest plant genera, Acacia, has undergone great change over the past 20 years and these changes have ramifications for weed biological control projects in a number of countries. Vachellia nilotica (prickly acacia) is a major weed in Australia, originating from the Indian subcontinent and Asia, and it has been a target for biological control since 1980. Once a member of Acacia, a large (>1,000 spp.) and iconic group in Australia, prickly acacia is now part of the genus Vachellia. Current knowledge suggests that Vachellia is more closely related to mimosoid genera than it is to Acacia s.s. There has also been a recent reclassification of legume subfamilies with subfamily Mimosoideae now part of subfamily Caesalpinioideae, and four new subfamilies. In this paper we review the changes that have occurred to this group since the prickly acacia biological control project began and discuss the implications for the project. A new host test list for quarantine testing is proposed. Developed following the modernisation of the centrifugal‐phylogenetic method, it is shorter than past lists, containing 46 species, although still lengthy because of the expectations of regulatory bodies, which are slower to accept advances in scientific knowledge. The list includes five Vachellia species, six “Mimoseae” species and 26 Acacia species. The number species from legume subfamilies other than the new Caesalpinioideae is greatly reduced.     

Tracking the movement of Nesidiocoris tenuis among banker plants and crops in a tomato greenhouse by DNA markers of host plants

The zoophytophagous predator, Nesidiocoris tenuis (Reuter) (Hemiptera: Miridae), is an important biological control agent. To maintain this insect, several non-crop host plants are used as banker plants in greenhouse crop systems. To optimize the efficiency of the predator-banker plant interaction, it is necessary to investigate how individual predators move between banker plants and crops. However, the movement is difficult to quantify under field conditions. Therefore, we investigated the movement of N. tenuis between tomato plants (Solanum lycopersicum L., Solanales: Solanaceae) and three banker plants (Cleome hassleriana Chod., Brassicales: Cleomaceae; Sesamum indicum L., Lamiales: Pedaliaceae; and Verbena × hybrida Voss, Lamiales: Verbenaceae) in a greenhouse by conducting PCR using plant-species-specific primers. Laboratory analysis results showed that our molecular method could detect N. tenuis activity within a relatively short time (≤ 24 h). In addition, N. tenuis predation on a pest species was unlikely to result in false detection of plant DNA in the pest (suggesting that N. tenuis had been on the plants). Multiple plant species were detected in adult insects collected from the greenhouse plants, indicating that N. tenuis frequently moved across the mentioned plant species. The movement patterns of N. tenuis between plant species varied substantially based on the plant species from which they were collected, which suggested each of the plant species had different functions for N. tenuis. Our findings revealed that planting multiple host plants would stabilize the N. tenuis population in biological control programs.

     

Chemical Constituents of Species in the Genus Pleione (Orchidaceae) and the Implications from Molecular Phylogeny

Pleiones are popular ornamental orchids and different species of Pleione are long being used as traditional medicine in many Asian countries. However, previous chemical investigations of the genus Pleione are restricted to only a few species. In the present study, high performance liquid chromatography (HPLC) fingerprint of Pleione plants was established, which in particular, eight common peaks were confirmed in 16 species/hybrids. Three of the compounds corresponding to the chromatographic peaks were identified by electrospray ionization tandem mass spectrometry (ESI-tandem-MS). HPLC analysis confirmed the studied taxa shared most of chemical compounds but the content of chemical compounds was significantly different between species. Comparison of hierarchical clustering result with phylogenetic tree revealed that closely related species have higher similarities in chemical constituents. In consideration of low chemical similarity between spring-flowering and autumn-flowering species, we suggest a discrimination of these two groups during medicinal use of the genus Pleione. Species with a large pseudobulb and with high content of a certain compound should be given priority in future artificial cultivation and medicinal cultivar breeding. We hope our findings will contribute to the quality control and promote conservation of such endangered plant group.     

Prioritisation of potential agents for the biological control of the invasive alien weed,Pereskia aculeata (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien species in South Africa that is native in Central and South America. In South Africa, P. aculeata outcompetes native plant species leading to a reduction in biodiversity at infested sites. Herbicidal and mechanical control of the plant is ineffective and unsustainable, so biological control is considered the only potential solution. Climatic matching and genotype matching indicated that the most appropriate regions in which to collect biological control agents were Santa Catarina and Rio de Janeiro provinces in Southern Brazil. Surveys throughout the native distribution resulted in 15 natural enemy species that were associated with the plant. Field host range data, as well as previous host plant records, were used to prioritise which of the species were most likely to be suitably host specific for release in South Africa. The mode of damage was used to determine which species were most likely to be damaging and effective if released. The most promising species prioritised for further study, including host specificity and impact studies, were the stem-wilter Catorhintha schaffneri Brailovsky & Garcia (Coreidae); the stem boring species Acanthodoxus machacalis Martins & Monné (Cerambycidae), Cryptorhynchus sp. (Curculionidae) and Maracayia chlorisalis (Walker) (Crambidae) and the fruit galler Asphondylia sp. (Cecidomyiidae). By prioritising the potential biological control agents that are most likely to be host-specific and damaging, the risk of conducting host specificity testing on unsuitable or ineffective biological control agents is reduced.     

Biological control of Rhamnus cathartica: is it feasible? A review of work done in 2002–2012

Rhamnus cathartica (common buckthorn) is a shrub (or small tree) of Eurasian origin, which has become invasive in North America. Internal feeders and sap suckers were prioritized for biological control from over 30 specialized insects identified from the target plant in its native European range. Five leaf‐feeding moths were also considered for further investigations. Field observations and preliminary host range tests with the stem‐boring beetle Oberea pedemontana, the root‐boring moth Synanthedon stomoxiformis, the shoot‐tip‐boring moth Sorhagenia janiszewskae and the leaf‐feeding moths Ancylis apicella, A. unculana, Triphosa dubitata, Philereme transversata and P. vetulata confirmed that all of these species were lacking host specificity in no‐choice conditions. Choice oviposition tests carried out with most of the prioritized species to assess their ecological host range yielded unreliable results. Three psyllids, Trichochermes walkeri, Cacopsylla rhamnicolla and Trioza rhamni are promising in terms of host specificity, but are infected with the plant disease ‘Candidatus Phytoplasma rhamni’. Fruit‐ or seed‐feeding insects may present the best potential for biological control of buckthorn in directly reducing seed set and thus seedling establishment. However, it was not possible to obtain adult fruiting trees of native North American Rhamnus species for testing. It is concluded that there are no promising arthropod agents based on what is known to date. Pathogens could offer new opportunities for biological control of R. cathartica in North America.     

Monophyletic clades of Macaranga‐pollinating thrips show high specificity to taxonomic sections of host plants

Thrips (Thysanoptera) have been recorded as pollinators of various plant species, but they are mostly regarded to be of low ecological relevance. In Southeast Asia, thrips were recently discovered to pollinate flowers of several taxonomic sections of the pioneer tree genus Macaranga (Euphorbiaceae), which is particularly well known as an ant‐plant, and for its importance in early forest succession. The lack of taxonomic treatment and of knowledge about systematic relationships among extant thrips, however, has prevented firm conclusions on the specificity of this plant–pollinator interaction. Here, results from sequencing of the mitochondrial cytochrome oxidase subunit support our previous morphospecies concept of Macaranga flower thrips, and confirm the genetic identity of five recently described species. They were remarkably all assigned to the genus Dolichothrips (Phlaeothripidae), which typically consists of phytophagous species. In addition, the molecular data revealed one cryptic species. A first phylogenetic tree of the Dolichothrips associated with Macaranga provides insights into their systematic position. In particular, we identify monophyly of all important Macaranga pollinator species, all species being largely specific to particular taxonomic host plant sections. Our results suggest a closely matched diversification of pollinating thrips with Macaranga trees. This adds a novel type of association to thrips pollinator–plant interactions, which have been so far documented as single‐species interactions or generalist thrips species visiting multiple plant taxa.     

Reverse influence of riparian buffer width on herbivorous and predatory Hemiptera

Insects are major contributors to farmland biodiversity, and their economic roles are also diverse. Many herbivorous species are crop pests, while predatory insects have the potential to act as biological controls against pests. Overall insect diversity has declined as a result of intensified agricultural practices. Riparian buffers may support insect populations in intensively cultivated areas, but their actual impact on the balance between harmful pests and beneficial predators is not known. It can be postulated that this impact may vary depending on the characteristics and location of the riparian buffer itself. We investigated the possibility that the biotic and abiotic attributes of agricultural riparian buffers adjoining crop fields and watercourses can explain the species composition of hemipteran assemblages. In particular, we were interested in the abundances of species belonging to the genus Nabis (generalist predators) and recognized and potential pests of cereal crops. Riparian buffer width and the presence or absence of woody plants were not associated with hemipteran species turnover among riparian buffers. In contrast, differences in the degree of dominance by grasses, in plant species turnover, and in which crop plant was cultivated in the adjacent field, explained a significant proportion of the variance in hemipteran species turnover. The abundance of predatory Nabis species increased with increasing riparian buffer width, whereas the abundance of recognized and potential crop pests decreased. The reverse patterns in the predatory and herbivorous Heteroptera suggest that increasing riparian buffer width might enhance biological control by Nabis predators.     

Response of predatory mites with different rearing histories to volatiles of uninfested plants

The behavioural response of the predatory mite Phytoseiulus persimilis to volatiles from several host plants of its prey, spider mites in the genus Tetranychus, was investigated in a Y-tube olfactometer. A positive response to volatiles from tomato leaves and Lima bean leaves was recorded, whereas no response was observed to volatiles from cucumber leaves, or leaves of Solanum luteum and Solanum dulcamara.Different results were obtained for predators that differed in rearing history. Predators that were reared on spider mites (Tetranychus urticae) on Lima bean leaves did respond to volatiles from Lima bean leaves, while predators that had been reared on the same spider mite species but with cucumber as host plant did not respond to Lima bean leaf volatiles. This effect is compared with the effect of rearing history on the response of P. persimilis to volatile allelochemicals of prey-infested plant leaves.     

Oviposition in Yponomeuta cagnagellus: the importance of contact cues for host plant acceptance

Small ermine moths (Yponomeutidae: Lepidoptera) are specialist herbivores. Species within the genus Yponomeuta are each specialized on a limited number of plant species, mainly within genera belonging to the Celastraceae. European Yponomeuta species have developed new specialized host affiliations, mainly on rosaceous hosts. Since these host shifts are reputed to be of consequence for speciation, the role of the ovipositing female is of particular interest. Study of the pre-oviposition behaviour of gravid Y. cagnagellus (Hb.) moths on host (Euonymus europaeus), non-host (Crateagus monogyna) and artificial oviposition substrates, provided information on the nature of the cues used for host plant acceptance and the insect’s perception of these cues. Host selection by adult females occurs with contact chemoreceptors probably located on the antennae or tarsi. MeOH-soluble, non-volatile phytochemical compounds washed from the host plant’s surface and applied on an artificial twig are sufficient to stimulate a complete sequence of behavioural elements leading to oviposition. Volatiles do not have a large effect on the pre-oviposition behaviour.     

Kanzawa spider mites acquire enemy‐free space on a detrimental host plant,oleander

Studies have proposed that predators of herbivores suffer significant fitness losses from the defense chemicals of host plants, and that herbivores adapted to these chemicals may experience reduced predation risk when residing on such plant species. We examined the effects of oleander, Nerium indicum Mill. (Apocynaceae), a host plant of the spider mite, Tetranychus kanzawai Kishida (Acari: Tetranychidae), on their prime predator, Neoseiulus womersleyi (Schicha) (Acari: Phytoseiidae), and tested the hypothesis that this host plant provides enemy‐free space. At the study sites, T. kanzawai occurred on oleander shrubs; in contrast, although N. womersleyi was present in the area, no individuals were found on oleander. Tetranychus kanzawai feeding on oleander negatively affected the settlement, development, and egg production of N. womersleyi. The lower egg production was a result of both the direct effects of oleander and the indirect effects via T. kanzawai. Previous studies showed that the fitness of T. kanzawai in the presence of N. womersleyi was lower than that in the absence of the predator, and lower on oleander than on other palatable host plant species in the absence of predators. Our findings suggest that N. womersleyi may not be able to invade T. kanzawai patches on oleander shrubs, which results in the fitness of T. kanzawai being higher on oleander than on other host plant species in the same area when N. womersleyi is present. This supports the hypothesis that T. kanzawai acquires enemy‐free space on oleander using the direct and indirect adverse effects of oleander on their predators as major defense mechanisms.     

Oviposition patterns and larval damage by the invasive horse‐chestnut leaf miner Cameraria ohridella on different species of Aesculus

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