Les bactéries rhizosphériques stimulatrices de croissance (PGPR) du genre Azospirillum sont utilisées commercialement pour leur capacité à stimuler la croissance et à augmenter le rendement des céréales via une relation associative complexe et peu comprise. L'objectif de cette étude a été d'utiliser les outils modernes de la métabolomique pour caractériser les mécanismes biochimiques activés par la souche A. lipoferum CRT1 chez son hôte, le maïs.L'analyse des contenus phytochimiques des racines, feuilles et sève ascendante a suggéré pour la première fois l'importance de la communication racine-feuille et des sucres simples dans l’augmentation de croissance et du potentiel de conversion photochimique de jeunes plantules par A. lipoferum CRT1. Une analyse transcriptomique a révélé un impact modéré au niveau des racines et des modifications de nombreux nœuds régulateurs des processus biologiques cellulaires des feuilles, dont ceux contrôlés par les auxines et l’acide abscissique. Des essais agronomiques conduits deux années consécutives sur quatre sites ont indiqué que l'augmentation de rendement par A. lipoferum CRT1 dérivait d’une sécurisation de la germination lors de stress environnementaux précoces et non de modifications des métabolomes (dont ceux liés à la nutrition azotée et phosphorée), de la croissance et du potentiel photosynthétique des plantules, des modifications qui étaient par contre liées aux contextes pédo-climatiques. La sécurisation de la germination était due à une accélération de la sortie de la radicule et de la consommation des sucres simples, et molécules apparentées, de la graine / Plant growth-promoting rhizobacteria (PGPR) of the genus Azospirillum are used commercially for their capacity to stimulate the growth and enhance the yield of cereal crops via an intricate, complex and poorly understood associative relationship. The aim of this study was to use modern metabolomics tools to decipher the biochemical mechanisms activated by A. lipoferum CRT1 in its maize host.The analysis of the phytochemical contents of roots, leaves and ascending sap revealed for the first time the importance of root-to-shoot communication and of simple sugars in the enhancement of growth and photochemical conversion potential of young plantlets by A. lipoferum CRT1. A transcriptomic analysis showed moderate impact on roots and a coordinate modulation of several regulatory nodes of cellular biological processes, including some mediated by auxins and abscissic acid. Agronomic field trials conducted two consecutive years in four sites correlated yield enhancement by A. lipoferum CRT1 to the securing of seed germination during environmental stresses. No correlation was found with modifications of plantlets metabolomes (including those linked to nitrogen and phosphorus nutrition), growth and photochemical conversion potential which were found to depend on additional soil and climate cues. Germination securing was due to a speeding of radicule emergence and of seed simple sugars consumption

Les maladies infectieuses sont une des causes les plus importantes de morbidité chez l'homme et l'animal avec des conséquences à la fois économiques, sanitaires et écologiques. L'étude de la diversité des pathogènes responsables et de leurs dynamiques de circulation au sein des communautés d'hôtes et de vecteurs, peut fournir des informations importantes pour la prévention et le contrôle de ces maladies. Dans ce contexte, nous nous sommes intéressés à l'agent pathogène responsable de la maladie de Lyme. Cette maladie est causée par les bactéries du complexe d'espèces Borrelia burgdorferi sensu lato (s.l.) transmises par les tiques lors de repas sanguins et sont capables d'infecter plusieurs espèces d'hôtes vertébrés. L'analyse de la diversité génétique de 63 souches de B. burgdorferi s.l., dont les génomes ont été séquencés, ont révélé un degré d'isolement génétique très important entre les différentes espèces du complexe. Les résultats obtenus suggèrent que les différents spectres d'hôtes des lignées de B. burgdorferi s.s. (principalement associées aux petits mammifères) et de B. garinii (normalement associées aux oiseaux) conduisent à des dynamiques de populations distinctes. De plus, grâce au séquençage haut-débit de deux marqueurs, nous avons pu démontrer qu'il existe, à une échelle intra-spécifique, des associations préférentielles des génotypes de B. burgdorferi avec différentes espèces de rongeurs. Enfin, en utilisant la diversité observée chez ces rongeurs et celle chez les tiques, nous avons estimé, via une approche de modélisation, que la contribution au risque de la maladie pour l'homme d'une espèce hôte introduite (tamia de Sibérie), pouvait être importante. / Infectious diseases are one of the major causes of human and animal morbidity, and they have impacts on the economy, public health, and the environment. By studying the diversity of the pathogens responsible for these diseases and their circulation within host communities and among vectors, we may glean valuable information that will aid prevention and control efforts. For these reasons, during my thesis, I became particularly interested in the pathogen(s) responsible for Lyme disease. This disease is caused by bacteria belonging to the Borrelia burgdorferi sensu lato (s.l.) species complex that are transmitted by ticks (during their blood meals) and that can infect several vertebrate host species. When I analyzed the genetic diversity present in 63 B. burgdorferi s.l. strains, whose genomes had been sequenced, I found that there was a significant degree of genetic separation among the different genospecies making up the complex. My results suggest that the fact that these different bacterial groups infect different ranges of hosts B. burgdorferi s.s. is mainly a pathogen of small mammals and B. garinii is primarily associated with birds lead to distinct population dynamics. Moreover, thanks to the high-throughput sequencing of two genetic markers, I have been able to show that, at an intraspecific level, certain B. burgdorferi genotypes are associated with specific rodent species. Finally, using the pathogen diversity observed in rodents and ticks, I employed a modeling approach to estimate the human disease risks presented by an introduced host species (the Siberian chipmunk) and found that these risks could be significant.

Avec l'intensification de la sylviculture et des échanges internationaux (introductions d'insectes ou plantations d'essences exotiques), les arbres forestiers sont de plus en plus confrontés à des attaques de nouveaux insectes phytophages. La compréhension des mécanismes d’adaptation de ces insectes à de nouveaux hôtes représente un enjeu scientifique majeur faisant intervenir de nombreux facteurs et des processus biologiques complexes. Dans ce contexte, nous nous sommes focalisés sur l’étude du rôle de deux facteurs : (1) la proximité taxonomique entre hôtes naturels et nouveaux de l'insecte ; (2) l’étendue du spectre d’hôtes naturels de l’insecte. Quatre approches complémentaires (méta-analyse, inventaire terrain, tests comportementaux, analyses génétiques), réalisées à différentes échelles taxonomiques et géographiques, ont été développées, les trois dernières approches se focalisant sur les relations scolytes indigènes - conifères introduits. Les objectifs étaient, d'une part d’approfondir les connaissances sur les capacités des insectes forestiers à étendre leur gamme d’hôtes, d'autre part d'apporter des informations utiles au gestionnaire forestier pour évaluer les risques économiques et écologiques qui découlent de ces adaptations. Les résultats des quatre approches concordent pour démontrer l’importance des deux facteurs testés dans les relations insectes-arbres en général et scolytesconifères en particulier. Les insectes à large spectre d’hôtes naturels seraient plus aptes à coloniser de nouveaux hôtes que ceux dont la gamme d’hôtes est plus restreinte. Parallèlement, la présence d'hôtes nouveaux proches taxonomiquement des hôtes naturels faciliterait le changement d’hôte. L’ensemble de nos résultats a contribué à compléter les connaissances sur l’évolution de la gamme d’hôtes des insectes forestiers phytophages en présence de nouveaux hôtes et par conséquent sur la sensibilité relative de ces hôtes par rapport aux hôtes autochtones. / With the intensification of forestry and global trade (insect introductions or plantations of exotic trees), forest trees have increasingly to face attacks by novel phytophagous insects. Understanding the adaptation mechanisms of these insects to their novel hosts, represents a crucial scientific issue involving many factors and complex biological processes. In this general topic, we focused on the role of two factors: (1) the taxonomic proximity between natural and novel hosts of the insects; (2) the host specificity of the insect. Four complementary approaches (meta-analysis, field inventory, behavioral tests and genetic analyses) at different taxonomic and geographical scales have been developed, the last three approaches focusing on the adaptation of indigenous bark beetles to exotic conifers. The objectives were, firstly to improve knowledge on the ability of forest insects to extend their natural host range, and secondly to provide useful information to forest managers, in order they can assess ecological and economic risks arising from these adaptations. Results from the four approaches agree to demonstrate the importance of both tested factors in the insect-tree relationships in general and conifer-bark beetle in particular. Forest insects with a broad natural host range appear to be better colonizers of novel hosts than insects with a narrow host range. The presence of novel tree species taxonomically related to native hosts seems to favor insect host shift. This work has also contributed to improve knowledge on the evolution of the host range of phytophagous forest insects in presence of novel host trees species, and consequently on the sensitivity of these hosts relatively to native ones.

La spécificité d'hôte est un concept fondamental pour comprendre les mécanismes évolutifs ayant abouti aux interactions étroites entre les bactéries et les plantes. Les études réalisées sur la coopération entre les bactéries rhizosphériques phytostimulatrices (PGPR) et les plantes suggèrent que la spécificité serait régie soit par une adaptation souchespécifique de la bactérie à des caractères aspécifiques de la plante, soit par une adaptation aspécifique de la bactérie à des propriétés spécifiques d'un génotype de plante. Ainsi, nous formulons l'hypothèse que ces adaptations se traduisent par la régulation de nombreux gènes indépendamment de leur implication directe dans l'effet phytobénéfique. Ces régulations pourraient dépendre de la combinaison souche bactérienne/génotype de plante considérée. L'objectif de ce travail est de mettre en évidence les gènes impliqués dans l'adaptation des partenaires l'un à l'autre et ceux impliqués dans la spécificité, au cours de la coopération PGPR-plantes. Pour cela, nous avons choisi comme modèle d'étude l'interaction entre Azospirillum lipoferum 4B et les céréales (blé, maïs, riz cultivars Cigalon et Nipponbare) avec un accent porté sur l'expression des gènes des deux partenaires de la coopération A. lipoferum 4B-riz. Les résultats de transcriptomique sur puce soulignent l'importance des mécanismes de réponse au stress oxydatif au cours de l'adaptation des partenaires l'un à l'autre ainsi que la mise en place de réseaux de régulation complexes. De nombreux gènes présentent un profil d'expression qui dépend de la combinaison souche/cultivar, suggérant que des mécanismes évolutifs ont conduit à une interaction préférentielle entre une souche et son cultivar d'origine / Host specificity is a fundamental concept in understanding evolutionary processes leading to intimate interactions between bacteria and plants. In the case of Plant Growth- Promoting Rhizobacteria (PGPR) specificity appears to be controlled either by a strainspecific bacterial adaptation to non-specific traits of the host plant or by non-specific bacterial adaptation to genotype-specific properties of the host plant. Thus, we hypothesize that these adaptations result in the regulation of a large number of genes, independently of their direct involvement in phytostimulation. These regulations may depend on the bacterial strain / plant genotype combination. This work aims at identifying genes involved in reciprocal adaptation of partners and those involved in host specificity in PGPR-plant cooperation. As a model, we studied the interaction between Azospirillum lipoferum 4B and cereals (wheat, corn, rice cultivars Nipponbare and Cigalon), with an emphasis on gene expression of both partners during A. lipoferum 4B-rice cooperation. Microarray transcriptomic results highlight the importance of mechanisms implicated in response to oxidative stress as well as a tight adjustment of regulatory networks during the adaptation of both partners to each other. Many genes display expression profile that depends on the strain/cultivar combination, suggesting that evolutionary processes have led to a preferential interaction between a strain and its original cultivar

Le contrôle biologique (C. -B. - l'utilisation d'ennemis naturels pour lutter contre les ravageurs) est durable, écologique et rentable pour contrer la résistance des ravageurs en augmentant l'utilisation des pesticides. Les parasitoïdes des pucerons sont des ennemis naturels communs des pucerons, les principaux ravageurs mondiaux dans l'agriculture. L'étude de la spécificité de l'hôte parasitoïde contribue à (1) comprendre les mécanismes écologiques et évolutifs de l'écosystème et (2) évaluer l'efficacité des agents de lutte biologique et les risques écologiques pour les espèces non ciblées. Cette étude porte sur la spécificité de l'hôte fondamental des parasitoïdes sur les niveaux individuels, en matière de besoins en ressources et dans le contexte des interactions multi trophiques sous stress abiotique environnemental, c'est-à-dire la limitation de l'eau. Aphidius ervi (Hymenoptera: Braconidés: Aphidiinae) a été choisi; ce parasitoïde puceron est largement utilisé comme modèle écologique et comme agent de lutte biologique commercial (BCA). D'une part, l'indice de spécificité de l'hôte A. ervi a été mesuré sur une large gamme d'espèces de pucerons. D'autre part, les impacts indirects de la limitation de l'eau ont été étudiés sur la spécificité de l'hôte du parasitoïde. En outre, les modifications induites par le stress hydrique dans la plante et les traits de vie des pucerons ont été mesurés. A. ervi s'est avéré être une espèce intermédiaire spécialisée qui a attaqué toutes les espèces de pucerons à des taux élevés, mais n'a pas pu se développer correctement sur toutes les espèces. Les quelques espèces qui se développaient bien étaient phylogénétiquement proches et appartenaient à la tribu des Macrosiphini. En outre, une corrélation positive de préférence - performance a été trouvée. Sous stress hydrique, la préférence et la performance des parasitoïdes ont été affectées, causant la perte de la corrélation. La limitation de l'eau a modifié négativement la qualité nutritionnelle de la plante, ce qui a entraîné une faible performance des pucerons sur les plantes hôtes. Ceci à son tour a diminué la convenance des hôtes pucerons pour le parasitoïde. Les effets de la limitation de l'eau n'étaient pas similaires pour toutes les combinaisons plantes-pucerons et dépendaient de plusieurs facteurs, à savoir les mécanismes végétaux adaptés au stress et la spécialisation de l'hôte des pucerons et des parasitoïdes. / Biological control (BC - the use of natural enemies to control pests) are sustainable, environmentally friendly and cost-effective methods to counteract pest resistance by increasing pesticide use. Aphid parasitoids are common natural enemies of aphids, the major worldwide pests in agriculture. The study of parasitoid host specificity contributes to (1) understanding ecological and evolutionary mechanisms driving the ecosystem and (2) evaluating the efficiency of biocontrol agents and the ecological risks for non-target species. This study focuses on the parasitoids fundamental host specificity on individual levels, in terms of resource requirements and in the context of multi-trophic interactions under environmental abiotic stress, i.e.water limitation. Aphidius ervi (Hymenoptera: Braconidae: Aphidiinae) was chosen; this aphid parasitoid is used widely as an ecological model and commercial biological control agent (BCA). On the one hand, A. ervi host specificity index was measured on a broad range of aphid species. On the other hand, the indirect impacts of water limitation were investigated on the host specificity of the parasitoid. Furthermore, water stress-induced modifications in the plant and the aphid life-history traits were measured. A. ervi was shown to be an intermediate specialist species who attacked all aphid species at high rates but was unable to develop well on all of them. The few that developed well were phylogenetically close and belong to the Macrosiphini tribe. Interestingly, a positive correlation preference – performance was found. Under water stress, both preference and performance of parasitoids were affected causing loss of the correlation. Water limitation negatively altered the plant nutritional quality resulting in low aphid performance on host plants. This in turn decreased the suitability of aphid hosts for the parasitoid. The impacts of water limitation were not similar across all plant-aphid combinations and depended on several factors, namely stress-adapted plant mechanisms and the host specialization of both aphids and parasitoids.

The Trypanosoma theileri group are blood parasites of ungulates. Confirmed vectors of these protists are horseflies or sheep keds for T. melophagium, but these trypanosomes were also detected in deer keds, mosquitoes and phlebotomus. We targeted on the investigation of possibility mosquitoes acting as vectors and additionally we targeted on the prevalence of T. theileri in mosquitoes at selected locations in the Czech Republic due to previous positive detection there. The measured prevalence has reached 6,4 % in mosquitoes, 4 % in deer keds and 16,7 % in sheep keds. The aim of experimental infections was to discover a potential of mosquitoes and phlebotomus acting as vectors of the T. theileri. Phlebotomus perniciosus and three species of mosquitoes Culex pipiens molestus, Culex pipiens quinquefasciatus and Aedes vexans were used for those experiments. Furthermore, isolates of T. theileri and T. melophagium from different sources were used. The divergence among these isolates and among used species of insect was found. The best results were achieved with Ae. aegypti and isolates from mosquitoes, where prevalence had reached 90,8 %. The infected mosquitoes were used for prediuresis research which had shown infection forms of the trypanosomes in the urine liquid. The phylogenetic analyse of T....

Lack of host-specific cell cultures necessitated initiation of primary cell cultures from ictalurid catfish. Cell cultures from the fin tissues of hybrid catfish ( channel catfish x blue catfish) were developed, characterized, and species of origin molecularly authenticated. Blue catfish alloherpesvirus (BCAHV) is an Ictalurid herpesvirus. When BCAHV was inoculated onto various fish cell lines from different families, the cytopathic effects were restricted to cell lines from family Ictaluridae indicating the host-specificity of virus. A virus challenge using channel, blue, and hybrid catfish indicated mortality due to BCAHV was significantly higher in blue and hybrid catfish. Crowding influenced BCAHV pathogenesis indicated by significantly higher mortality in highly stocked tanks. Host susceptibility to BCAHV differed with age. Temperature had a significant role in the activation and pathogenesis of BCAHV. The enhanced virulence of BCAHV in blue and hybrid catfish reveals its potential to be a significant pathogen in catfish culture.

Trygonorrhina fasciata (Rhinobatidae) specimens naturally infected by three monogenean species were captured and maintained in marine aquaria to promote a continuous parasite load. Monogenean eggs recovered from aquaria provided larvae for descriptions and life history experiments. I describe the adult, larva and post-larval development of a new species of hexabothriid, Branchotenthes octohamatus, from the gills. This is the first monogenean larva described with only eight hooklets. This character may be useful to help resolve problematic relationships within the Hexabothriidae and offers insight into more general hypotheses about relationships within the Monogenea. I also redescribe the adult of Calicotyle australis (Monocotylidae) from the cloaca and describe the larva. The number and arrangement of larval ciliated epidermal cells and sensilla was revealed using silver nitrate. I redescribe Pseudoleptobothrium aptychotremae (Microbothriidae) adults from the skin of T. fasciata, representing a new host and locality record. Larval anatomy and post-larval development are also documented. The presence of six needle-like spicules in the larval haptor is confirmed, supporting an earlier theory that spicules are ancestral vestiges. My studies revealed three different egg hatching, host finding strategies and larval ‘types’. Branchotenthes octohamatus has a ‘sit-and-wait’ strategy, entirely dependent on mechanical disturbance to stimulate eggs to hatch. Larvae are unciliated, cannot swim, lack pigmented eyespots and show no photo-response but may survive for more than two days after hatching at 22ºC. In contrast, eggs of C. australis hatch spontaneously with a strong diurnal rhythm in the first few hours of daylight when exposed to a LD12:12 illumination regime. Larvae are ciliated and can swim, have pigmented eyespots, are photo-positive and can remain active and survive for up to 24 h after hatching at 22ºC. Eggs of P. aptychotremae may have a ‘bet-hedging’ strategy. Some eggs hatch spontaneously and rhythmically in an LD12:12 regime during the last few hours of daylight but their low hatching success rate suggests that other eggs may require a different cue provided by the host. Larvae are ciliated, can swim, lack pigmented eyespots, show no photo-response and remain active for only a few hours at 22ºC. Experiments using the fluorescent dye, 5(6)-carboxyfluorescein diacetate N-succinimidyl ester (CFSE) revealed B. octohamatus on gills of T. fasciata within 30 min of exposure to the host. This provides strong evidence that larvae invade the gills directly via the host’s inhalant respiratory current and do not migrate after initial attachment elsewhere. Five rhinobatid species (Aptychotrema vincentiana, T. fasciata, Trygonorrhina sp. A, A. rostrata and Rhinobatos typus), with overlapping distributions spanning west, south and east Australian coastal waters were surveyed for monogeneans at four locations between Fremantle, Western Australia and Stradbroke Island, Queensland. Genetic homogeneity, using the mitochrondrial gene Cytochrome b (cytb) and the nuclear marker, Elongation factor-1 alpha (EF1a), was observed for all Branchotenthes and Calicotyle specimens irrespective of collection locality or rhinobatid species. Genetic homogeneity was observed for Pseudoleptobothrium specimens collected in western and southern Australia. However, local genetic heterogeneity was apparent among Pseudoleptobothrium specimens collected from two sympatric host species in New South Wales. Analyses revealed a highly divergent clade, indicating a morphologically cryptic, ancestral species. / http://proxy.library.adelaide.edu.au/login?url= http://library.adelaide.edu.au/cgi-bin/Pwebrecon.cgi?BBID=1323070 / Thesis (Ph.D.) -- University of Adelaide, School of Earth and Environmental Sciences, 2008

Trygonorrhina fasciata (Rhinobatidae) specimens naturally infected by three monogenean species were captured and maintained in marine aquaria to promote a continuous parasite load. Monogenean eggs recovered from aquaria provided larvae for descriptions and life history experiments. I describe the adult, larva and post-larval development of a new species of hexabothriid, Branchotenthes octohamatus, from the gills. This is the first monogenean larva described with only eight hooklets. This character may be useful to help resolve problematic relationships within the Hexabothriidae and offers insight into more general hypotheses about relationships within the Monogenea. I also redescribe the adult of Calicotyle australis (Monocotylidae) from the cloaca and describe the larva. The number and arrangement of larval ciliated epidermal cells and sensilla was revealed using silver nitrate. I redescribe Pseudoleptobothrium aptychotremae (Microbothriidae) adults from the skin of T. fasciata, representing a new host and locality record. Larval anatomy and post-larval development are also documented. The presence of six needle-like spicules in the larval haptor is confirmed, supporting an earlier theory that spicules are ancestral vestiges. My studies revealed three different egg hatching, host finding strategies and larval ‘types’. Branchotenthes octohamatus has a ‘sit-and-wait’ strategy, entirely dependent on mechanical disturbance to stimulate eggs to hatch. Larvae are unciliated, cannot swim, lack pigmented eyespots and show no photo-response but may survive for more than two days after hatching at 22ºC. In contrast, eggs of C. australis hatch spontaneously with a strong diurnal rhythm in the first few hours of daylight when exposed to a LD12:12 illumination regime. Larvae are ciliated and can swim, have pigmented eyespots, are photo-positive and can remain active and survive for up to 24 h after hatching at 22ºC. Eggs of P. aptychotremae may have a ‘bet-hedging’ strategy. Some eggs hatch spontaneously and rhythmically in an LD12:12 regime during the last few hours of daylight but their low hatching success rate suggests that other eggs may require a different cue provided by the host. Larvae are ciliated, can swim, lack pigmented eyespots, show no photo-response and remain active for only a few hours at 22ºC. Experiments using the fluorescent dye, 5(6)-carboxyfluorescein diacetate N-succinimidyl ester (CFSE) revealed B. octohamatus on gills of T. fasciata within 30 min of exposure to the host. This provides strong evidence that larvae invade the gills directly via the host’s inhalant respiratory current and do not migrate after initial attachment elsewhere. Five rhinobatid species (Aptychotrema vincentiana, T. fasciata, Trygonorrhina sp. A, A. rostrata and Rhinobatos typus), with overlapping distributions spanning west, south and east Australian coastal waters were surveyed for monogeneans at four locations between Fremantle, Western Australia and Stradbroke Island, Queensland. Genetic homogeneity, using the mitochrondrial gene Cytochrome b (cytb) and the nuclear marker, Elongation factor-1 alpha (EF1a), was observed for all Branchotenthes and Calicotyle specimens irrespective of collection locality or rhinobatid species. Genetic homogeneity was observed for Pseudoleptobothrium specimens collected in western and southern Australia. However, local genetic heterogeneity was apparent among Pseudoleptobothrium specimens collected from two sympatric host species in New South Wales. Analyses revealed a highly divergent clade, indicating a morphologically cryptic, ancestral species. / http://proxy.library.adelaide.edu.au/login?url= http://library.adelaide.edu.au/cgi-bin/Pwebrecon.cgi?BBID=1323070 / Thesis (Ph.D.) -- University of Adelaide, School of Earth and Environmental Sciences, 2008

Host specificity of tropical bark and ambrosia beetles was surveyed by rearing the beetles from 13 host trees in a lowland rainforest in Papua New Guinea. Ploeophagous bark beetles show narrow host specificity (usually family-level) typical for herbivorous insects, fungus-growing ambrosia beetles display almost no host fidelity. In both groups of species, the local diversity of plants is unlikely to have played a role in the clade diversification. The ambrosia symbiosis (scolytine beetles and fungi) is shown to be less specific than previously assumed, based on a discovery of new association between Scolytodes unipunctatus (genus of phloem feeders) and three unrelated groups of ambrosia fungi. The hypothesis that apparent polyphagy may conceal specialized populations within a species of a herbivore is tested for Homona mermerodes (Lepidoptera, Tortricidae). The haplotype diversity of the species show no congruence with host plants or geographic origin, confirming polyphagy of the species.