P.J. De Barro

Refined global analysis of Bemisia tabaci (Hemiptera: Sternorrhyncha: Aleyrodoidea: Aleyrodidae) mitochondrial cytochrome oxidase 1 to identify species level genetic boundaries.

ABSTRACT Identifying species boundaries within morphologically indistinguishable cryptic species complexes is often contentious. For the whitefly Bemisia tabaci (Gennadius) (Hemiptera: Sternorrhyncha: Aleyrodoidea: Aleyrodidae), the lack of a clear understanding about the genetic limits of the numerous genetic groups and biotypes so far identified has resulted in a lack of consistency in the application of the terms, the approaches used to apply them and in our understanding of what genetic structure within B. tabaci means. Our response has been to use mitochondrial gene cytochrome oxidase one to consider how to clearly and consistently define genetic separation. Using Bayesian phylogenetic analysis and analysis of sequence pairwise divergence we found a considerably higher number of genetic groups than had been previously determined with two breaks in the distribution, one at 11% and another at 3.5%. At >11% divergence, 11 distinct groups were resolved, whereas at >3.5% divergence 24 groups were identified. Consensus sequences for each of these groups were determined and were shown to be useful in the correct assignment of sequences of unknown origin. The 3.5% divergence bound is consistent with species level separations in other insect taxa and suggests that B. tabaci is a cryptic species composed of at least 24 distinct species. We further show that the placement of Bemesia atriplex (Froggatt) within the B. tabaci in group adds further weight to the argument for species level separation within B. tabaci. This new analysis, which constructs consensus sequences and uses these as a standard against which unknown sequences can be compared, provides for the first time a consistent means of identifying the genetic bounds of each species with a high degree of certainty.

Asymmetric Mating Interactions Drive Widespread Invasion and Displacement in a Whitefly

The role of behavioral mechanisms in animal invasions is poorly understood. We show that asymmetric mating interactions between closely related but previously allopatric genetic groups of the whitefly Bemisia tabaci, a haplodiploid species, have been a driving force contributing to widespread invasion and displacement by alien populations. We conducted long-term field surveys, caged population experiments, and detailed behavioral observations in Zhejiang, China, and Queensland, Australia, to investigate the invasion process and its underlying behavioral mechanisms. During invasion and displacement, we found increased frequency of copulation leading to increased production of female progeny among the invader, as well as reduced copulation and female production in the indigenous genetic groups. Such asymmetric mating interactions may be critical to determining the capacity of a haplodiploid invader and the consequences for its closely related indigenous organisms.

Reproductive incompatibility among genetic groups of Bemisia tabaci supports the proposition that the whitefly is a cryptic species complex

The worldwide distribution and extensive genetic diversity of the whitefly Bemisia tabaci has long been recognized. However, whether B. tabaci is a complex species or a species complex has been a subject of debate. Recent phylogenetic analyses suggest that B. tabaci is a cryptic species complex composed of at least 24 morphologically indistinguishable species. Here, we conducted crossing experiments and demonstrated reproductive incompatibility among three of the 24 putative species. Our data and those of previously reported crossing experiments among various putative species of B. tabaci were collated to reveal the pattern of reproductive isolation. The combined results provide strong support to the proposition that B. tabaci is a cryptic species complex.

Bemisia argentifolii is a race of B. tabaci (Hemiptera: Aleyrodidae): the molecular genetic differentiation of B. tabaci populations around the world

The phylogenetic relationships between genotypes of Bemisia tabaci were compared using ITS1 and CO1 nucleotide sequences. Phylogenetic and minimum spanning network analyses identified six major races, Asia, Bali, Australia, sub-Saharan Africa, Mediterranean/Asia Minor/Africa and New World as well as a large collection of genotypes from the Asia region with no strong association with any of the races. The term race is based on its usage in Mallet (2001). Mating incompatibility occurs between some races. There is insufficient data to raise races to species status, but the data supports the recognition of the six races and an unresolved core of ungrouped genotypes under the single Bemisia tabaci (Gennadius) species name. To clarify the identity of the race to which the B. tabaci under investigation is known, the following nomenclature is suggested, B. tabaci (Asia), B. tabaci (Bali), B. tabaci (Australia) B. tabaci (sub-Saharan Africa), B. tabaci (Mediterranean/Asia Minor/Africa) and B. tabaci (New World). Further, there is insufficient molecular or biological data to support the separation of B. tabaci and B. argentifolii Bellows & Perring and its use should be discontinued.

Genetic structure of the whitefly Bemisia tabaci in the Asia–Pacific region revealed using microsatellite markers

Bemisia tabaci (Hemiptera: Aleyrodidae) is a haplo‐diploid species of sap‐feeding insect belonging to the group of insects commonly known as whiteflies. From earlier analyses of mitochondrial and ribosomal markers it has been concluded that in the Asia–Pacific region there were three major indigenous races as well as a large collection of genotypes with no clear association with any race. This new study uses 15 microsatellite loci and demonstrates that the indigenous Asia–Pacific genotypes can be split into six genetic populations with little or no gene flow between them. These bare only superficial similarity to the mitochondrial and ribosomal defined races. Moreover, four of the six can be further split into two subpopulations that again show little evidence gene flow between them. While the patterns reflect a strong geographical structure, physical barriers alone cannot explain all the observed structure. Differential host‐plant utilization explained some of the substructure, but could not explain the overall structure. The roles of mating interference and Wolbachia in developing the genetic structure are considered. The lack of gene flow between genetic populations and some subpopulations further suggests that the barriers were either sufficiently impermeable to immigration or that reproductive isolation and competitive interactions were sufficiently strong to prevent gene flow. If the latter is the case, it suggests that there may be as many as 10 morphologically indistinguishable species indigenous to the Asia–Pacific region.

Host Plant and Biotype Density Interactions – Their Role in the Establishment of the Invasive B Biotype of Bemisia tabaci

Bemisia tabaciis a complex of closely related genetic types of whiteflies, few of which are invasive. One of these, B biotype, has proven to be particularly adapted to invading new areas, but the underlying reasons as to why it has a well-developed capacity to invade is not known. To develop an understanding of factors that may be contributing to B’s invasive capacity, inter-biotype mating interactions and host plant suitability for the exotic B (B. tabaci Mediterranean/Asia Minor/Africa) and the indigenous Australian (AN) biotype (B. tabaci Australia) were examined. The results suggest that when confined to a mutually acceptable host, B cannot establish when the ratio of AN : B exceeds 20 : 1. However, when simultaneously provided with a host that only it prefers, B is able to establish even at 50 : 1 (AN : B). Further, when both biotypes occur together the number of progeny per female increases (relative to the number produced when only one biotype is present). The response is observed for both biotypes, but is considerably greater in the case of B. In addition, B performs better in the presence of the AN biotype B. tabaci Australia while AN perform worse in coexistence with B, but only if the demographics allow B to mate without significant interference. This leads to the prediction that B will invade in circumstances where its unique hosts are of sufficient number to escape the full negative impact of inter-biotype mating interactions and reduced competitiveness in terms of reproductive rate, while exposing the indigenous biotype to the full effects of the interaction.

Species within the Bemisia tabaci (Hemiptera: Aleyrodidae) Complex in Soybean and Bean Crops in Argentina

ABSTRACT The whitefly Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) is a cryptic species complex that contains some of the most damaging pests in tropical and subtropical regions. Recent studies suggested that this complex is composed of at least 24 distinct species. We use the approach from these studies to consider the identity of B. tabaci in Argentina. Previous studies have suggested the presence of a B. tabaci presumably indigenous to the Americas and referred to as the BR biotype in Argentina. We placed the entity referred to as the BR biotype within the B. tabaci cryptic species complex using whiteflies collected in soybean and bean crops in northern and central Argentina. The whiteflies were assigned using the mitochondrial cytochrome oxidase (mtCOI) gene. Four unknown haplotypes plus two Argentina sequences from GenBank formed a cluster that was basal to the rest of the New World sequences. These sequences diverged from the consensus sequence across the range of 3.6 to 4.3%. Applying the species assignment rules of recent studies suggests that the individuals from Argentina form a separate species. A fifth unknown haplotype fell within the New World putative species and formed a distinct cluster with haplotypes from Panama. These results suggest that Argentina has two indigenous species belonging to the B. tabaci cryptic species complex. Rather than using mtCOI sequencing for all B. tabaci collected, a simple random amplified polymorphic DNA-polymerase chain reaction diagnostic was used and tested along with previously published primers designed to work specifically with the BR biotype from Brazil. These primers were either unable to distinguish between the two indigenous members of the complex in Argentina or indicated a difference when none was evident on the basis of mtCOI sequence comparison.

Further insights into the strange role of bacterial endosymbionts in whitefly, Bemisia tabaci : Comparison of secondary symbionts from biotypes B and Q in China

The percentage infection of secondary symbionts (SS) (Wolbachia, Arsenophonus, Rickettsia, Hamiltonella, Fritschea and Cardinium) in the exotic Bemisia tabaci (Genn.) invaders, commonly known as biotypes B and Q from China, were determined by PCR. In total, 373 biotype B and 1830 biotype Q individuals were screened for the presence of SS. Biotype B was more abundant than biotype Q from 2005 to 2006, and biotype Q was more abundant from 2007 to 2009. Each of the SS, with the exception of Fritschea, was detected in both biotypes B and Q; Fritschea was found in none of the samples examined. For biotype B, the percentage infection of Hamiltonella was the highest (92.0%) followed by Rickettsia (70.2%). For biotype Q, the percentage infection of Hamiltonella was again the highest (73.3%). Arsenophonus was the least common of the SS observed in both biotypes B and Q. The percentage infection of Wolbachia, Rickettsia and Hamiltonella in biotype B was each significantly higher than in biotype Q, whereas the percentage infection of Cardinium in biotype B was significantly lower than in biotype Q. The percentage infection of SS in biotypes B and Q varied from year to year over the period 2005-2009. Furthermore, within biotype Q, two distinct subgroups were identified which differ from each other in terms of their SS complement. We discuss these results in the light of the potentially influential factors and roles of the SS.

The biology of two Eretmocerus spp. (Haldeman) and three Encarsia spp. Forster and their potential as biological control agents of Bemisia tabaci biotype B in Australia

The performance, as measured by daily rate of parasitism and total parasitism, of five aphelinid species found in Australia parasitising Bemisia tabaci were compared on cotton, hibiscus, rockmelon, soybean and tomato. Two Eretmocerus spp., both indigenous to Australia, gave the highest levels of parasitism on each of the plant host species tested. The tritrophic interactions between B. tabaci, host plant species and Eret. mundus (Australian parthenogenetic form) (APF) were also examined. In general, more whiteflies were parasitised when cotton was the source host or rockmelon the test host. Parasitism was always low when tomato was either the source or test host. When parasitoids were transferred from rockmelon to cotton, parasitism declined. In contrast, parasitism increased when parasitoids were transferred from cotton to rockmelon. Parasitism also increased when parasitoids were transferred from soybean to rockmelon, yet failed to do so when shifted from soybean to cotton despite cotton normally being a better host. However, when parasitoids were transferred from cotton to soybean there was a marked increase in parasitism. Possible causes are discussed. The field cage trial demonstrated that parasitism by both Eretmocerus spp. increased with increasing whitefly density. Further, the increase in parasitism was not due to the presence of more parasitoids as neither the parasitoid‐whitefly ratio nor the total number of parasitoids present had a significant effect on parasitism. The combination of the two species, gave similar levels of parasitism to that achieved by Eret. mundus (APF) alone. Subsequent identification of the emerged individuals indicated that over 50% of the parasitism was due to this species suggesting that it out‐competed Eret. queenslandensis. Despite this competition, there was no evidence that overall control was compromised.

The role of temperature, photoperiod, crowding and plant quality on the production of alate viviparous females of the bird cherry‐oat aphid, Rhopalosiphum padi

Experiments indicated that for offspring of apterous Rhopalosiphum padi (L.) (Hemiptera: Aphididae), photoperiod and crowding were the most important determinants of wing development whereas crowding and plant quality were more significant for the next generation. Plant quality became increasingly important as temperature increased while crowding became less so. More alates developed on plants previously infested with aphids, indicating that aphid feeding reduced plant quality. High temperature suppressed alatoid production, but could be overcome by crowding. Temperature appeared to influence wing development indirectly rather than directly by acting on the aphid through the plant. Adult weight and potential fecundity were also reduced for aphids which fed on previously infested plants.