Presence of Cicadomorpha in olive orchards of Greece with special reference to Xylella fastidiosa vectors

Insects of the infraorder Cicadomorpha cause serious economic impact on agriculture because of pathogens’ transmission. The vector-borne bacterium Xylella fastidiosa, which is associated with olive quick decline syndrome, is exclusively transmitted by insects of this infraorder and more specific by sharpshooters and spittlebugs. Efficient control practices of the insect vectors are necessary for the control of the disease in cases of outbreaks. Therefore, the knowledge of the vectors' species as well as the robust understanding of their biology and ecology is of crucial importance. In the present study, extensive sampling was conducted in 28 olive orchards of Greece during the years 2017 and 2018 aiming to investigate which species of this infraorder are present, with special interest to those which are considered as vectors of the bacterium. During the surveys, 4,350 Cicadomorpha insects were collected. Forty-two, five and one species of the families Cicadellidae, Aphrophoridae and Cercopidae, respectively, were identified. The results of the surveys revealed that the main potential vectors of X. fastidiosa in Greek olive orchards are the spittlebugs Philaenus spumarius and Neophilaenus campestris. Those species have wide distribution in Greece and are present for a long period during the year, and they inhabit both the ground vegetation and the olive trees' canopy. A representative portion of the captured spittlebugs was analysed with molecular methods, and the analysis confirmed the absence of X. fastidiosa in the tested individuals.     

Introducing cultivated trees into the wild: Wood pigeons as dispersers of domestic olive seeds

Animals may disperse cultivated trees outside the agricultural land, favoring the naturalization or, even, the invasiveness of domestic plants. However, the ecological and conservation implications of new or unexplored mutualisms between cultivated trees and wild animals are still far from clear. Here, we examine the possible role of an expanding and, locally, overabundant pigeon species (Columba palumbus) as an effective disperser of domestic olive trees (Olea europaea), a widespread cultivated tree, considered a naturalized and invasive species in many areas of the world. By analyzing crop and gizzard content we found that olive fruits were an important food item for pigeons in late winter and spring. A proportion of 40.3% pigeons consumed olive seeds, with an average consumption of 7.8 seeds per pigeon and day. Additionally, most seed sizes (up to 0.7 g) passed undamaged through the gut and were dispersed from cultivated olive orchards to areas covered by protected Mediterranean vegetation, recording minimal dispersal distances of 1.8–7.4 km. Greenhouse experiments showed that seeds dispersed by pigeons significantly favored the germination and establishment in comparison to non-ingested seeds. The ability of pigeons to effectively disperse domestic olive seeds may facilitate the introduction of cultivated olive trees into natural systems, including highly-protected wild olive woodlands. We recommend harvesting ornamental olive trees to reduce both pigeon overpopulation and the spread of artificially selected trees into the natural environment.     

Dispersal capacity of Monochamus galloprovincialis,the European vector of the pine wood nematode,on flight mills

The pine wood nematode (Bursaphelenchus xylophilus), which causes the symptoms of pine wilt disease, is recognized worldwide as a major forest pest. It was introduced into Portugal in 1999. It is transmitted between trees almost exclusively by longhorn beetles of the genus Monochamus, including, in particular, M. galloprovincialis (Coleoptera: Cerambycidae) in maritime pine forests. Accurate estimates of the flight capacity of this insect vector are required if we are to understand and predict the spread of pine wilt disease in Europe. Using computer‐linked flight mills, we evaluated the distance flown, the flight probability and speed of M. galloprovincialis throughout adulthood and investigated the effects of age, sex and body weight on these flight performances, which are proxies for dispersal capacity. The within‐population variability of flight performance in M. galloprovincialis was high, with a mean distance of 16 km flown over the lifetime of the beetle. Age and body weight had a significant positive effect on flight capacity, but there was no difference in performance between males and females. These findings have important implications for managing the spread of the pine wood nematode in European forests.     

Transmission of Xylella fastidiosa by naturally infected Philaenus spumarius (Hemiptera,Aphrophoridae) to different host plants

The recent establishment of Xylella fastidiosa subspecies pauca in the southern Italian region of Apulia threatens agricultural crops and the environment. Olive is an important and widespread ancient crop in Italy and, so far, the most impacted host. The meadow spittlebug Philaenus spumarius (Hemiptera, Aphrophoridae) has been identified as a vector of X. fastidiosa in southern Italy; this species is one of the most common potential vectors in Europe. To generate disease management strategies, data on X. fastidiosa transmission by P. spumarius are necessary. Therefore, we carried out transmission experiments by using field‐collected spittlebugs in 2014 and 2015 (5 and 11 collection dates, respectively), and transferring groups of insects immediately on to recipient plants. Various host plant species were tested: olive, oleander, sweet orange, grapevine and the stone fruit rootstock GF677 (Prunus persica × Prunus amygdalus). Xylella fastidiosa was detected in all the host plants after insect plant access except for grapevine; infections to sweet orange and stone fruit were not systemic. In 2015, estimates of insect X. fastidiosa infectivity were obtained; the number of PCR‐positive P. spumarius on each plant was positively correlated with the plant infection status. The proportion of P. spumarius infected with X. fastidiosa ranged from 25% to 71% during the entire survey period. The number of X. fastidiosa cells detected in P. spumarius heads ranged from 3.5 × 10 to 4.0 × 10² (CFU equivalents), which is lower than that reported for leafhopper vectors in the Americas. These data show that field‐collected P. spumarius have high rates of X. fastidiosa infection and are competent vectors.     

Interrelation of mating,flight, and fecundity in navel orangeworm females

The navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae, Phycitini), is an economically important pest of nut crops in California, USA. Improved management will require better understanding of insect dispersal, particularly relative to when mating occurs. A previous study demonstrated a more robust laboratory flight capacity compared to other orchard moth pests, but it was unclear how mating affects dispersal, and how dispersal affects fecundity. In this study, 1‐ and 2‐day‐old females were allowed to fly overnight on a flight mill either before or after mating, respectively, and were then allowed to oviposit. Data on fecundity were compared between treatments to minimally handled or tethered‐only control females. Females that mated before flight flew longer and covered a greater distance than those flying prior to mating. However, timing of flight relative to mating did not affect fecundity, nor did any measure of flight performance. There was no effect on fecundity when females were forced to fly for designated durations from 3 min to 2 h. Together, our data revealed no obvious trade‐off between flight activity and reproductive output. Distances measured on the flight mills (mean ca. 15 km for mated females) may overestimate net displacement in the field where flight tracks are often meandering. The results suggest that most females mate and oviposit in or near their natal habitat, but that some may disperse potentially long distances to oviposit elsewhere.     

Cabbage whiteflies colonise Brassica vegetables primarily from distant,upwind source habitats

The occurrence of species in rapidly changing environments, such as agricultural landscapes, is affected by their ability to recolonise habitats. Knowledge of the landscape scale affecting colonisation is essential for large‐scale pest management. Colonisation by insects can be affected on multiple landscape scales, as different morphs of a species may have specific dispersal abilities. The cabbage whitefly, Aleyrodes proletella (L.) (Hemiptera: Aleyrodidae), a major pest of Brassica vegetables, is known to colonise Brassica vegetables primarily from fields of oilseed rape, Brassica napus L. (Brassicaceae). We used field mapping and remote sensing to characterise the relevant scales for colonisation of Brussels sprouts by cabbage whiteflies. Surprisingly, oilseed rape fields in wide landscapes (2–8 km around study sites) explained colonisation better than oilseed rape areas in local landscapes (200–1 000 m around study sites). The explained variance increased when additional weight was given to upwind source habitats, indicating wind transport of whitefly colonisers. Low importance of local compared to wide landscape source habitats can be explained by the flight behaviour of whitefly morphs. Migratory morphs show phototactic attraction but are attracted by hosts only during the later phases of flight. Therefore, they ignore host plants close to their origin and disperse several kilometres. Trivial flight morphs rarely move more than a few hundred metres. In conclusion, as most whitefly colonisers reached Brassica vegetables from source habitats at a distance of 2–8 km, predictions on pest pressure and landscape‐scale whitefly management should consider these distances. In contrast, oilseed rape fields in the local landscape, which usually worry farmers, had little effect.     

Seasonal flight activity by the Asian citrus psyllid in east central Florida

The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), is an important invasive citrus pest in the USA because it vectors a bacterium responsible for huanglongbing, a devastating disease of citrus. Information was lacking on seasonal aspects of flight activity by D. citri, which could have ramifications on psyllid management as well as our understanding of epidemiology of the disease. Of interest from a pest management standpoint would be whether D. citri regularly disperses to or away from citrus on a predictable schedule. In research presented here, seasonal flight activity by D. citri was investigated using yellow sticky traps deployed in citrus trees and in fallow areas adjacent to citrus. Results indicated that flight activity by both male and female D. citri away from citrus can occur at any time of the year with consistent dispersal activity during the spring. The research further indicated citrus is continually subject to infestation by immigrating adults and that there is no time during the year that a citrus grower could be assured immigration would not occur. Growers should be aware that adult dispersal occurs regularly during spring and they should time management tactics accordingly. Adult flight activity 2 m from a citrus tree was more pronounced at 1 m above ground than at 2 or 3 m high. At distances of 8–60 m from trees, numbers of adults on traps were similar among the three heights. Males and females were similar with respect to seasonal flight activity. Numbers of adults captured on traps distant from citrus were not correlated with wind speed, sunlight, or air temperature, but there was some evidence that relative humidity influenced flight activity. Although the D. citri life cycle is dependent on flush, data from these studies did not confirm that psyllid dispersal from citrus consistently increases as citrus flush abundance decreases.     

Parallel host shifts in a bacterial plant pathogen suggest independent genetic solutions

While there are documented host shifts in many bacterial plant pathogens, the genetic foundation of host shifts is largely unknown. Xylella fastidiosa is a bacterial pathogen found in over 600 host plant species. Two parallel host shifts occurred—in Brazil and Italy—in which X. fastidiosa adapted to infect olive trees, whereas related strains infected coffee. Using 10 novel whole-genome sequences from an olive-infecting population in Brazil, we investigated whether these olive-infecting strains diverged from closely related coffee-infecting strains. Several single-nucleotide polymorphisms, many derived from recombination events, and gene gain and loss events separated olive-infecting strains from coffee-infecting strains in this clade. The olive-specific variation suggests that this event was a host jump with genetic isolation between coffee- and olive-infecting X. fastidiosa populations. Next, we investigated the hypothesis of genetic convergence in the host shift from coffee to olive in both populations (Brazil and Italy). Each clade had multiple mutations and gene gain and loss events unique to olive, yet no overlap between clades. Using a genome-wide association study technique, we did not find any plausible candidates for convergence. Overall, this work suggests that the two populations adapted to infect olive trees through independent genetic solutions.     

Dispersal of the leafhoppers <Emphasis Type="Italic">Matsumuratettix hiroglyphicus</Emphasis> and <Emphasis Type="Italic">Yamatotettix flavovittatus</Emphasis> (Homoptera: Cicadellidae), vectors of sugarcane white leaf disease

Sugarcane white leaf disease is a serious problem in many Asian Countries. The leafhoppers Matsumuratettix hiroglyphicus (Matsumura) and Yamatotettix flavovittatus Matsumura are the main vectors of sugarcane white leaf disease phytoplasma. Gaining a better understanding of the dispersal behavior of such insect vectors is essential for both disease epidemiology and vector control. The dispersal distances of M. hiroglyphicus and Y. flavovittatus in a sugarcane field were estimated by means of mark–release–recapture experiments. Adult leafhopper vectors collected from the fields were marked using fluorescent dye powder and released at a central release point in a sugarcane field. The marking method did not have a significant effect on the survival or flight activity of the leafhopper vectors. The overall release–recapture rates of M. hiroglyphicus and Y. flavovittatus within 50 m were 10.1 and 13.4 %, respectively. The estimated natural mean dispersal distances for M. hiroglyphicus and Y. flavovittatus were 162.1 and 387.5 m, respectively. Wind appears to be the main factor influencing the leafhopper dispersal direction.     

Significance of habitat type for the genetic population structure of Panonychus citri (Acari: Tetranychidae)

Restricted migration and habitat fragmentation promote genetic differentiation between populations: Because most of the hosts of Panonychus citri are woody plants, mainly citrus trees that are usually planted at intervals of several metres, this mite likely faces more risks (e.g., starvation) by dispersing between host plants, compared to other spider mite species that infest both herbaceous and woody plants, such as Tetranychus urticae. Such a limited gene flow between patches (host plants) can lead to differentiation of populations even within a small area. Therefore, we hypothesize that P. citri populations are genetically differentiated not only between distant populations but also within small areas, such as within a grove. To test this hypothesis, we investigated the divergence of P. citri populations in Japanese citrus groves according to a hierarchical arrangement of geographical distance, ranging from distant populations (10 groves distributed throughout different areas in two major Japanese islands; this level of analysis is referred to as ‘geographic’) to local populations (different trees in a specific grove; ‘local’). Three molecular markers were used an esterase locus, one microsatellite and a point mutation in the mitochondrial cytochrome oxidase subunit I. At a local level acaricide susceptibility tests were also performed using two acaricides: fenpyroximate (25 ppm) and etoxazole (3.33 ppm). At a broad geographic level the gene diversity decreased with decreasing area size and distance between populations. By contrast, at the local level, populations maintained a significant level of variation between trees within groves, and the divergence within groves was higher than between groves. Whereas no statistical difference of the mortalities was detected among groves for the two acaricides tested, the difference was statistically significant among trees within groves in fenpyroximate (ANOVA, p < 0.025) and=" marginal=" in=" etoxazole="><>p < 0.05). we=" concluded=" that=">P. citri populations maintain a higher level of variation between trees (or patches of trees) within groves than between groves at the local level, though the gene diversity tended to be smaller with decreasing distance between populations at the geographical level. Results are discussed in relation to the dispersal behaviour of spider mites.     

Dispersal differences of a pest and a protected Cerambyx species (Coleoptera: Cerambycidae) in oak open woodlands: a mark–recapture comparative study

1. Cerambyx welensii (Cw) and Cerambyx cerdo (Cc) are two large saproxylic beetles living on Quercus trees in the Western Palearctic whose current pest and legal status differs markedly. Cw is an emerging pest involved in oak decline while Cc is an internationally protected species. 2. Acquiring knowledge of the ecology, demography and behaviour of Cw and Cc in forests harbouring mixed populations is a demanding task to optimise their management or protection. Here, we report the results of a mark–recapture study on the flight behaviour and dispersal potential of both species in holm oak open woodlands. 3. Average flights, dispersal rates and diffusion models were remarkably similar in Cw and Cc, reflecting a low‐dispersal tendency and sedentary behaviour. However, a subset of adults in both species exhibited a huge propensity to disperse and flew more than 1 km. An aggregated distribution of dispersal distances suggested that a behavioural polymorphism might underlie the flight pattern. 4. Adults moved preferentially in those spatial directions with neighbouring trees, while prevailing winds did not significantly affect dispersal patterns. The main interspecific differences were as follows: (i) Cc performed longer crosswind flights than Cw on windy days; (ii) Cc tended to fly farther than Cw at low temperatures; and (iii) adult feeding improved dispersal but only in small Cw females. Moon phase did not affect flight activity. 5. The results are discussed in an effort to interpret how these ecological and behavioural differences might shape the life history of both congeneric species when they live in sympatry in dehesa woodlands.     

Artificial diet delivery system for Philaenus spumarius,the European vector of Xylella fastidiosa

Artificial diets represent an essential tool for investigations on the intimate relationship between plant pathogens and their vectors. Previous research failed in devising an artificial diet delivery system for the meadow spittlebug Philaenus spumarius, to date considered the most important vector of the bacterium Xylella fastidiosa in Europe. Here, we describe a new delivery “tube system” by which we succeeded in artificial feeding of P. spumarius with holidic diets (one sucrose diet and two amino acids diets). Spittlebug probing and feeding behaviour on either the tube system or a traditional “flat system” realized out of a small Petri dish filled with diet and covered with stretched Parafilm^® were observed in real time by video‐EPG (Electrical Penetration Graph), in order to assess the occurrence of ingestion and excretion. Moreover, we evaluated P. spumarius survival on either the tube system filled with the two holidic diets that gave the best EPG results or an empty tube system serving as control. Contrary to the flat system, where just brief stylet insertions through the Parafilm^® were recorded, the spittlebug ingested the artificial diets when delivered with the tube system. Survival on the diets provided with the tube system was significantly greater than the control, with no differences between the diets tested. Furthermore, the tube system was suitable also for another spittlebug species shown to be a competent vector of X. fastidiosa, that is Neophilaenus campestris. The tool we devised opens new perspectives for investigations on X. fastidiosa/spittlebugs interactions, as well as for the functional analysis of mutant X. fastidiosa strains in respect to insect colonization and transmission.     

Population structure of Guyanagaster necrorhizus supports termite dispersal for this enigmatic fungus

Understanding how species accomplish dispersal of their propagules can shed light on how they are adapted for their ecosystem. Guyanagaster necrorhizus is a sequestrate fungus, meaning its dispersal propagules, or spores, are entirely enclosed within a fruiting body, termed a sporocarp. This fungus is most closely related to Armillaria and its allies. While Armillaria species form mushrooms and have forcibly discharged spores, G. necrorhizus spores have lost this ability, and by necessity, must be passively dispersed. However, G. necrorhizus does not possess characteristics of other sequestrate fungi with known dispersal mechanisms. Repeated observations of termites feeding on G. necrorhizus sporocarps, and spores subsequently adhering to their exoskeletons, led to the hypothesis that termites disperse G. necrorhizus spores. To test this hypothesis, we used microsatellite markers and population genetics analyses to understand patterns of clonality and population structure of G. necrorhizus. While Armillaria individuals can spread vegetatively over large areas, high genotypic diversity in G. necrorhizus populations suggests spores are the primary mode of dispersal. Spatial genetic structure analyses show that G. necrorhizus sporocarps within 238 m of each other are more closely related than would be expected by chance and conservative estimates from population assignment tests suggest gene flow no longer occurs between sporocarps separated by 2 km. These distances are consistent with previous studies analysing foraging distances of the termites found associated with G. necrorhizus sporocarps. Termites have rarely been recorded to specifically target fungal sporocarps, making this a potentially novel fungal–insect interaction.     

Hunter feedback of individually marked wild boar <Emphasis Type="Italic">Sus scrofa</Emphasis> L.: dispersal and efficiency of hunting in northeastern Germany

Increasing wild boar (Sus scrofa L.) population densities all over Europe cause severe economic problems. For understanding mechanisms of epidemics, the knowledge of dispersal is required. Thus, we investigated dispersal rates and distances with regard to sex and age of wild boar in southwestern Mecklenburg-Western Pomerania. From 152 marked wild boar, 105 have been registered as dead, of which, 51% were males and 49% females. Forty-five percent were shot as piglets, 41% as yearlings, and 14% as adults. The distance between capture site and site of death ranged between 184 m and 41.5 km. Piglets were shot closer to their capture site (mean distance 1 km) than older animals (mean 4 km), although this difference was only significant for males. In general, males tended to disperse further before being shot (3.8 km) than females (1.6 km). Only 3.8% of all animals were shot at distances larger than 10 km. As most animals (84.6%) were shot inside their natal home range, only a small proportion (15.4%) did actually disperse (shot outside mothers home range), which is 32% of all animals surviving to the age of yearlings. Of those dispersed animals, 25% were females. The low dispersal rate is biased by female philopatry and allows actual dispersal only at very high population densities or in sparsely populated regions. In consideration for the low natural mortality proved by radio-tagged animals, the harvest rate is lower than the net reproduction. We did not detect any sex-biased hunting. The dominating hunting method was single hunt at bait, although drive hunts are highly effective. However, hunting rates on piglets and females were too low for regulating the population.     

Dispersal and dormancy strategies among insect species competing for a pulsed resource

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Estimation of bumblebee queen dispersal distances using sibship reconstruction method

Dispersal ability is a key determinant of the propensity of an organism to cope with habitat fragmentation and climate change. Here we quantify queen dispersal in two common bumblebee species in an arable landscape. Dispersal was measured by taking DNA samples from workers in the spring and summer, and from queens in the following spring, at 14 sites across a landscape. The queens captured in the spring must be full sisters of workers that were foraging in the previous year. A range of sibship reconstruction methods were compared using simulated data sets including or no genotyping errors. The program Colony gave the most accurate reconstruction and was used for our analysis of queen dispersal. Comparison of queen dispersion with worker foraging distances was used to take into account an expected low level of false identification of sister pairs which might otherwise lead to overestimates of dispersal. Our data show that Bombus pascuorum and B. lapidarius queens can disperse by at least 3 and 5 km, respectively. These estimates are consistent with inferences drawn from studies of population structuring in common and rare bumblebee species, and suggest that regular gene flow over several kilometres due to queen dispersal are likely to be sufficient to maintain genetic cohesion of ubiquitous species over large spatial scales whereas rare bumblebee species appear unable to regularly disperse over distances greater than 10 km. Our results have clear implications for conservation strategies for this important pollinator group, particularly when attempting to conserve fragmented populations.     

Recruitment of two Opuntia species invading abandoned olive groves

In Europe, many agricultural areas are now abandoned and hence can be invaded by exotic species. The abundance and spatial distribution patterns of two Opuntia species were studied in old olive groves in the Parc Natural del Cap de Creus, Catalonia (Spain). Seedling recruitment (97.3% and 51.5% of juveniles for O. maxima and O. stricta, respectively) was higher than recruitment by cladodes. O. maxima had more seedlings recruited beneath olive trees and beneath Opuntia adults than expected. Most O. stricta seedlings were also located beneath Opuntia adult plants. However, although most seedlings were recruited beneath Opuntia, some (10–30%) were found away from putative parental plants. This may be due to seed dispersal by birds and wild boars. Seeds dispersed by wild boars were not significantly more viable than seeds from intact fruits. Seedlings grow very slowly but have a high survival rate. In conclusion, Opuntia seedling recruitment is very successful and ensures the persistence of these species within old olive groves. Consequently, it prevents restoration from an agricultural land-use back to the native community.     

Potential vectors of Xylella fastidiosa: a study of leafhoppers and treehoppers in citrus agroecosystems affected by Citrus Variegated Chlorosis

This study investigated the predominant leafhopper and treehopper (Hemiptera, Auchenorrhyncha) species in Citrus Variegated Chlorosis (CVC)‐affected citrus agroecosystems in Argentina, their seasonal fluctuation, and their potential role as vectors of Xylella fastidiosa Wells et al., using molecular methods for detection. More than 6 000 Auchenorrhyncha were collected from three citrus agroecosystems over a period of 3 years using yellow sticky traps and entomological nets. Cicadellidae and Membracidae were the most abundant families. Of the 43 species identified, five were predominant in citrus orchards, and three were predominant in weeds surrounding citrus plants. All predominant species and another four non‐predominant species tested positive for X. fastidiosa in PCR and real‐time PCR assays. In a transmission assay, Dechacona missionum (Berg), Tapajosa rubromarginata (Signoret), and Cyphonia clavigera (Fabricius) transmitted X. fastidiosa successfully. Scaphytopius bolivianus Oman and Frequenamia spiniventris (Linnavuori) populations increased once (during the summer), possibly due to favorable weather conditions, and Bucephalogonia xanthophis (Berg), Molomea lineiceps Young, and T. rubromarginata populations increased twice a year: once in summer and once in winter, coinciding with the increase in early citrus shoots (flush). Among the X. fastidiosa‐positive species, those with the higher population densities during the sprouting period, where trees are highly susceptible to infection, must be considered as most relevant vectors of CVC in the citrus‐growing areas in Argentina.     

Morphological variation associated with dispersal capacity in a tree‐killing bark beetle Dendroctonus ponderosae Hopkins

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Two different Xylella fastidiosa strains circulating in Italy: phylogenetic and evolutionary analyses

Xylella fastidiosa, a bacterial species infecting a broad range plants, includes five subspecies, fastidiosa, multiplex, pauca, mulberry and sandyi. In Europe, Xylella was isolated in olive trees in southern Italy (Apulia region) during the year 2013. The aim of the present study was to apply phylogenetic and evolutionary analysis to trace the possible origin and way of the entrance of Xylella fastidiosa in Italy. All the genomes available for Xylella fastidiosa spp were downloaded from NCBI. A phylogeographic analysis was performed using BEAST. X. fastidiosa strains belonging to X. fastidiosa subsp. pauca and subsp. sandyi have been reported to infect olive trees and coffee plants, respectively. The phylogeographic analysis also revealed and confirmed these two different ways of provenience X. fastidiosa subsp. pauca from Costa Rica and X. fastidiosa subsp sandyi from California Phylogeny have been an important tool to validate and support the recent hypothesis for X. fastidiosa pauca provenience.