Prevalence of a new genetic group, MEAM-K, of the whitefly Bemisia tabaci (Hemiptera: Aleyrodidae) in Karnataka, India, as evident from mtCOI sequences.
The nymphs and adults actively feed on the phloem sap and excrete copious amounts of honeydew, which supports the growth of sooty mold that in turn impedes photosynthesis (Byrne &Bellows 1991). Several studies examined the host-related phenotypic variations in B. tabaci and concluded that variations are evident not only among the populations on different hosts but also among individuals occurring on the same host (Mound 1963; Palaniswami et al. 1996; Lisha et al. 2003). Morphometric analysis of 4th instar nymphs revealed phenotypic variations corresponding to variations in the leaf anatomy of the host plant (Maruthi et al. 2007). Bemisia tabaci is known to be an aggressive colonizer of crops, with varying traits at the morphological (Bellows et al. 1994; Costa et al. 1995; Rosell et al. 1997), biochemical (Costa &Brown 1991; Brown et al. 2000; Perring 2001), and molecular levels (Gawel & Bartlett 1993; De Barro et al. 2005; Boykin et al. 2007).
Different geographical and ecological variations with host plant specialization offer an ideal system for the study of sympatric speciation in B. tabaci, direct damage to crops and vector ability (Chu et al. 2007). De Barro et al. (2011) stated that asymmetrical mating interference provides a clear mechanism that could contribute to formation of global genetic structure. Genetic differentiation of different populations in the species complex was analyzed mainly based on the ribosomal internal transcribed spacer 1 (rITS-1) and mitochondrial cytochrome oxidase I (mtCOI) sequences worldwide (Simon et al. 1994; Hu et al. 2011; Zasada et al. 2014). Dinsdale et al. (2010) identified 3.5% pair-wise genetic divergence as the considered boundary for separating different species; this was further supported with either complete or partial mating isolation between a number of putative B. tabaci "species" (Xu et al. 2010; Wang et al. 2011). Considering all the past taxonomical approaches, Boykin et al. (2012) defined species in the B. tabaci species complex. In the present study, we examined the genetic variation in the mtCOI region of the genome of B. tabaci, collected on 30 different host plants from different locations in Karnataka, India, to assess the presence and prevalence of various genetic groups.
Materials and Methods
Nymphs and adults of B. tabaci were collected with a handheld aspirator from different locations in Karnataka, India, on 30 different hosts (Fig. 1 and Table 1) and preserved in 70% alcohol until further use.
GENOMIC DNA EXTRACTION AND AMPLIFICATION
The total DNA was extracted from individual adult B. tabaci specimens using DNAeasy Tissue Kit (Qiagen GmbH, Hilden, Germany) according to the manufacturer's protocol and stored at -80[degrees]C until further use. Polymerase chain reaction (PCR) was performed with mtCOI primers F-C1-J-2195 (5'-TTGATTTTTTGGTCATCCAGAAGT-3') and R-L2N-3014 (5'-TCCAATGCACTAATCTGCCATATTA-3') (Simon et al. 1994; Frohlich et al. 1999). The 25 [micro]L PCR consisted of 50 ng/[micro]L template,10 picomoles of each primer, 0.25 mM dNTP mix, 1.5 mM Mg[Cl.sub.2], 1 U Taq polymerase, and 2.5 [micro]L Taq buffer (Fermentas GmBH, St. Leon-Rot, Germany). The PCR cycling was carried out in a thermal cycler (Applied Biosystems, Veriti 96 wells, USA) with the following parameters: 94[degrees]C for 5 min as initial denaturation followed by 35 cycles of denaturation at 94[degrees]C for 30 s, annealing at 54[degrees]C for 45 s, and extension at 72[degrees]C for 50 s, and 72[degrees]C for 10 min as the final extension. The amplified products were resolved in 1.0% agarose gel, stained with ethidium bromide (10 Hg/mL), and visualized under UV light. Amplified PCR products were eluted using the gel extraction kit Nucleospin[R] Extract II (Macherey-Nagel, Duren, Germany) according to the manufacturer's protocol, and sequencing was performed in M13 forward and reverse directions.
All mtCOI sequences corresponding to different genetic groups of B. tabaci were downloaded from the National Center for Biotechnology Information (NCBI) GenBank (Suppl. Table 1, available online at http://purl.fcla.edu/fcla/entomologist/browse). Sequence alignment was performed employing MUSCLE implemented in Seaview (Thompson et al. 1994). Genetic divergence was calculated employing MEGA5 (Tamura et al. 2011). The nucleotide substitution model for the best fits and the model parameters were estimated using Akaike Information Criterion implemented in the program MODELTEST 3.7 (Posada &Crandall 1998) in conjunction with PAUP*. Maximum parsimony and maximum likelihood analyses were performed with PAUP* 4.0b10 (Swofford 1998), using the heuristic search procedure with 1,000 random additions of sequences and 10 trees held at each pseudo-rep licate, and the tree bisection reconnection branch swapping method with all characters was treated as unordered and equally weighted. The chosen model with estimated parameters was used to derive the maximum likelihood tree in RaxML with the heuristic search settings. The same software was used to generate consensus trees using the CONSENSE program (http://evolution.genetics.washington.edu/phylip/doc/consense.html), and the tree was rooted with the outgroup Bemisia afer (Priesner & Hosny) (Hemiptera: Aleyrodidae). Phylogenetic analysis was performed using MrBayes (Huelsenbeck & Ronquist 2001). MrBayes 3.1 was run for 10 million generations by using 8 chains and sampled every 1,000 generations. All runs reached a plateau in the likelihood score, and the same was indicated by standard deviations of split frequencies (0.0023). Our 4 Markov Chain Monte-Carlo chains were converged, indicated by the potential scale reduction factor, which was close to one. The burn-in parameter was estimated employing Tracer version 1.5 (Rambaut & Drummond 2009), and the trees corresponding to the first 20% of generations were discarded. We performed 2 independent MrBayes runs to ensure the analyses were not trapped in the local optima, after which topologies and posterior probabilities (PP) from these 2 runs were compared for congruence purpose. The trees obtained were visualized using FigTree v1.3.1 (Rambaut 2009).
The mtCOI gene was successfully amplified and sequenced from 71 individual adult B. tabaci. We analyzed 131 sequences, including the 71 generated from this study. The sequence analysis revealed that out of 816 bp, 332 nucleotides were conserved, 484 were variable, and 379 were parsimony informative. Absence of stop codons indicated that no pseudogenes were amplified within the sequences, and similar base composition indicated no indels. Bemisia tabaci nucleotide frequencies were 24.46% (A), 43.10% (T/U), 19.23% (C), and 13.22% (G). Rates of different transitional substitutions are shown in bold and those of transversional substitutions are shown in italics in Table 2. The base composition of the mtCOI gene fragment was biased toward adenine (A) and thymine (T) with an overall 67.56%. The transition/transversion rate ratios were [k.sub.1] = 6.255 (purines) and [k.sub.2] = 6.190 (pyrimidines). The overall transition/transversion bias was R = 2.629. All sequences were deposited in the NCBI GenBank with the accession numbers KF790634 to KF790689 and HQ268814, HQ268813, HQ268812, HQ268811, HQ331246, HQ331245, and HQ331244.
Based on the phylogenetic analyses, the Indian B. tabaci samples were clustered into Asia-I, Asia-II-7, Asia-II-8, Middle East Asia Minor-1 (MEAM-1), and MEAM-K groups (Figs. 2a and 2b). The Asia-I genetic group was predominant representing 44 of the 71 (61.97%) samples sequenced. The Asia-I population had 17 different host plant species (Table 1), of which Solanum lycopersicum L., Solanum melongena L. (Solanales: Solanaceae), Gossypium hirsutum L., and Abelmoschus esculentus (L.) Moench (Malvales: Malvaceae) were the most frequent host plants and are widely distributed in Karnataka (Fig. 2b). Few samples collected from Karnataka clustered with AsiaII-7 and were found on 9 host plant species. The Asia-II-8 genetic group contained 5 samples, which had been collected on 5 different host plants. The most important genetic groups MEAM-1 and MEAM-2 formed 2 subclusters representing B and B2 genotypes. Here, MEAM-1 comprised 5 sequences, which had been collected on 3 host plants (mostly cabbage). However, 3 sequences from specimens collected on Phaseolus vulgaris L. (Fabales: Fabaceae) at Kolar, H-Cross, and Malur (Karnataka, India) diverged from all the MEAM-1, MEAM-2, Indian Ocean, and Mediterranean groups. Thus, they formed an independent genetic group, which we named MEAM-K group. The genetic divergence ranged from 4.0 to 24.0% with an average of 15.7% (Table 3 and Fig. 3).
"There is probably no other concept in biology that has remained as consistently controversial as the species concept" (Mayr 1982). This quote is remarkably true with B. tabaci, because this species is difficult to be diagnosed using morphological, cytological, behavioral, molecular, and biochemical methods, and its identification is complicated further by reproductive isolation (Rosell et al. 1997; Maruthi et al. 2007). Molecular studies on various insects of agricultural importance have helped to identify new species (Ball &Armstrong 2006), biotypes (Perring 2001), cryptic species (Hebert et al. 2004), and haplotypes (Toda &Murai 2007), which are difficult to identify through morphology due to phenotypic plasticity and lack of distinguishing morphological features (Russell 1957; Mound 1963; Rosell et al. 1997; Maruthi et al. 2007).
Considering the vector potential of B. tabaci, it is necessary to analyze the molecular diversity of the same species or species complex collected on various host plants. In this regard, the resistant tomato varieties Arka Ananya (against tomato leaf curl virus) and Arka Abhay and Arka Anamika (against Bhendi yellow vein mosaic virus) have become susceptible again, and this breakdown of resistance may be correlated with the existence of various species of vectors or virus, but the exact plant physiological mechanism is still unknown. In 1991, the biotype nomenclature was introduced to the B. tabaci species complex based on esterase banding patterns but was no longer used after the advent of Random Amplified Polymorphic DNA-PCR and mtCOI and ITS sequencing (Perring 2001; Simon et al. 2003; Zang et al. 2006; Boykin 2013). In the discussion on B. tabaci nomenclature, it remains unclear whether to call a new biotype a genetic group or a putative species with novel binomial nomenclature, such as Bemisia argentifolii Bellows & Perring (De Barro et al. 2011; Boykin et al. 2013).
Bemisia tabaci can attack a wide range of host plants globally, and its species complex is composed of at least 34 morphologically indistinguishable species (Boykin et al. 2012, 2013; Boykin 2014). Different cropping patterns and diverse climatic conditions may be responsible for the apparent diversity in B. tabaci, which otherwise is grouped by geographic location (Rekha et al. 2005). The current study revealed the existence of 5 genetic groups within Karnataka State, India, namely Asia-I, Asia-II-7, Asia-II-8, MEAM-1, and MEAM-K, with the latter being a new group identified in this study.
The genetic divergence ranged from 4.0 to 24.0% with an average of 15.7%. According to Dinsdale et al. (2010), the genetic divergence among 198 mtCOI sequences of B. tabaci and sequences of the outgroup species B. afer, B. atriplex (Froggatt), and B. subdecipiens Martin (Hemiptera: Aleyrodidae) ranged from zero to 34%. However, Lee et al. (2013) analyzed the genetic divergence after excluding the outgroup sequences and concluded that it ranged from zero to 24.0%, which was on par with our study.
Of the reported 5 genetic groups, the major group was Asia-I with members collected on 19 different host plants although most had been collected on eggplant; this group was found previously to be prevalent across Asia (Boykin et al. 2007; Dinsdale et al. 2010; Hu et al. 2011). The eggplant-associated Asia-I B. tabaci can transmit several begomo-viruses (Govindappa 2002; Muniyappa et al. 2003) and was shown to transmit eggplant yellow mosaic virus in Thailand (Green et al. 2003). In this study, we report another 17 host plants for Asia-I including cotton, pumpkin, ridge gourd, okra, capsicum, sunflower, potato, carrot, mustard, and tomato. However, a previous study showed that tomato is not a preferred host plant for Asia-I because it is associated with low fecundity (Chowda-Reddy et al. 2012). Host-associated variations in B. tabaci influence its rate of fecundity, which could be due to premating or post-mating selection against migrants and hybrid progeny (Liou & Price 1994; Brunner et al. 2004). However, the most to least preferred hosts, in terms of oviposition, were eggplant, cotton, pumpkin, tomato, and cassava (Venkatesh 2000). Our study also supported this finding, wherein eggplant and cotton were the most frequent host plants for Asia-I. Apart from the Jatropha genetic group, which occurs in a separate ecological niche, most of the putative species exhibit high levels of polyphagy (Burban et al. 1992; Brown et al. 1995). According to Chowda-Reddy et al. (2012), MEAM-1 has a wide host range and produces higher quantities of honeydew than other genetic groups. This is also true for genetic groups Asia-I, Asia-II-7, and Asia-II-8, of which Asia-I had the greatest number of host plants in our study.
We identified Asia-II-7 from Karnataka occurring on 9 different host plants, mostly on ornamental plants. Asia-II-7 was reported first in 1998 in India (Ramappa et al. 1998) and in China (Qiu et al. 2006). A previous study suggested that this putative species adapts readily to ornamental plants rather than vegetables (Shah et al. 2013). The 3rd genetic group identified in this study was Asia-II-8, which was recently named as B. gossypiperda (Boykin 2014) and was collected on 5 different host plants including cotton (Chowda-Reddy et al. 2012). Significantly, we here report the occurrence of a previously unreported subclade for Middle East genetic groups (MEAM-1 and 2). We named it MEAM-K because the samples were collected on P. vulgaris from Kolar, Karnataka, the native location for MEAM-1. For Kolar and nearby areas where both species have become established, our study raises questions about the relative pest status of MEAM-1and MEAM-K. We do not know to what extent these genetic groups differ with respect to important factors such as their ability in feeding and reproduction, their efficiency as vectors, or their susceptibility to insecticides. These high levels of molecular and ecological resolution will be needed to mitigate quarantine disputes arising from the detection of morphologically indistinguishable members of the B. tabaci genetic groups. Thus, our study may stand as a link to detect and identify B. tabaci genetic groups, determine areas of occurrence, identify areas of invasion, and design management strategies.
In conclusion, the current study revealed the existence of 5 genetic groups of B. tabaci in Karnataka, India, identified as Asia-I, Asia-II-7, Asia-II-8, MEAM-1, and a previously unreported genetic group, MEAMK. Thus, our work will help in rapid and accurate identification of these putative genetic groups of B. tabaci, which in turn will help in further elucidating the epidemiology and management of geminiviruses and be of value in the operation of quarantines.
This paper is part of the doctoral degree work of the senior author. We gratefully acknowledge the financial support received for the "Outreach Programme on Management of Sucking Pests in Horticultural Crops" from the Indian Council for Agricultural Research (ICAR), New Delhi, India. We are thankful to the director at the Indian Institute of Horticultural Research (IIHR) for the support and facilities.
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H. K. Roopa (1), *, R. Asokan (1), K. B. Rebijith (1), Ranjitha H. Hande (1), Riaz Mahmood (2), and N. K. Krishna Kumar (3)
(1) Division of Biotechnology, Indian Institute of Horticultural Research, Hessaraghatta Lake P.O Karnataka, Bengaluru 560 089, India
(2) Department of Biotechnology and Bioinformatics, Kuvempu University, Jnanasahyadri, Shankaraghatta, Shimoga577 451, India
(3) Division of Horticulture, ICAR, Krishi Anusandhan Bhawan - II, New Delhi 110 012, India
* Corresponding author; E-mail: firstname.lastname@example.org
Supplementary material for this article in Florida Entomologist 98(4) (Dec 2015) is online at http://purl.fcla.edu/fcla/entomologist/browse
Caption: Fig. 1. Map indicating different locations of Bemisia tabaci sampling and distribution of genetic groups in Karnataka, India.
Caption: Fig. 2a. Phylogenetic tree showing the relationship of the Bemisia tabacimt mtCOI sequences collected in this study to consensus sequences of Dinsdale et al. (2010). Bayesian analyses were performed employing MrBayes under the best-fit model GTR+I+G of molecular evolution for 20 million generations and 25% discarded as burn-in. Posterior probabilities are shown above the branches and maximum likelihood scores from RaxML indicated below the branches. Asia-I condensed is shown in Fig. 2b.
Caption: Fig. 2b. Phylogenetic tree indicating the Asia-I group, which was the most abundant genetic group in this study.
Caption: Fig. 3. Graph indicating net evolutionary divergence between groups of sequences.
Table 1. Details of Bemisia tabaci mtCOI sequences generated for this study along with place of collection, accession number, voucher specimen number, respective genetic group, and host plants. SI no. Host Common name 1 Ocimum sanctum Tulsi 2 Parthenium hysterophorus Aster (white top weed) 3 Raphanus sativus Radish 4 Unknown Ornamental Ornamental 5 Manihot esculenta Tapioca 6 Hibiscus cultivar Hibiscus 7 Gossypium hirsutum Cotton 8 Cucumis sativus Cucumber 9 Solanum melongena Eggplant 10 Brassica oleraceo Cabbage 11 Musa acuminata Banana 12 Phaseolus vulgaris Common beans 13 Brassica oleraceo Cabbage 14 Brassica oleraceo Cabbage 15 Lablab purpureus Hyacintha beans 16 Abelmoschus esculentum Okra 17 Citrullus la not us Watermelon 18 Lycopersicon seculentum Tomato 19 Gossypium hirsutum Cotton 20 Lagenaria sice rana Bottle gourd 21 Lycopersicon esculentum Tomato 22 Musa acuminta Banana 23 Luffa spp. Ridge gourd 24 Abelmoschus esculentus Okra 25 Helia nth us annuus Sunflower 26 Capsicum annuum Capsicum 27 Cucurbita moschata Pumpkin 28 Vigna unguiculata Cowpea 29 Citrullus lanatus Watermelon 30 Daucus carota Carrot 31 Brassica juncea Mustard 32 Crossandra infundibuliformis Crossandra 33 Cucumis sativus Cucumber 34 Gossypium hirsutum Cotton 35 Solanum melongena Eggplant 36 Ricinus communis Castor bean 37 Cucumis melo Muskmelon 38 Gossypium hirsutum Cotton 39 Solanum tuberosum Potato 40 Brassica rapa Turnip 41 Solanum melongena Eggplant 42 Solanum melongena Eggplant 43 Solanum melongena Eggplant 44 Solanum melongena Eggplant 45 Solanum melongena Eggplant 46 Abelmoschus esculentum Bhendi 47 Solanum melongena Eggplant 48 Solanum melongena Eggplant 49 Solanum melongena Eggplant 50 Solanum melongena Eggplant 51 Solanum tuberosum Potato 52 Cucumis sativus Cucumber 53 Abelmoschus esculentum Bhendi 54 Lycopersicon esculentum Tomato 55 Gossypium hirsutum Cotton 56 Gossypium hirsutum Cotton 57 Solanum melongena Eggplant 58 Gossypium hirsutum Cotton 59 Solanum melongena Eggplant 60 Gossypium hirsutum Cotton 62 Brassica oleraceo Cabbage 63 Gossypium hirsutum Cotton 64 Solanum melongena Eggplant 65 Gossypium hirsutum Cotton 66 Solanum melongena Eggplant 67 Lycopersicon esculentum Tomato 68 Gossypium hirsutum Cotton 69 Solanum melongena Eggplant 70 Phaseolus vulgaris Beans 71 Phaseolus vulgaris Beans 72 Phaseolus vulgaris Beans Bangalore, Accession SI no. Karnataka, India number Latitude Longitude 1 Malleshwarum KF790634 12.98 77.57 2 Tarabanahalli KF790635 13.11 77.47 3 IIHR KF790636 13.13 77.48 4 Malleshwarum KF790637 12.98 77.57 5 Malleshwarum KF790638 12.98 77.57 6 Malleshwarum KF790639 12.98 77.57 7 Hessaraghatta KF790653 13.13 77.48 8 Andaman KF790667 11.68 92.77 9 Chamarajanagar KF790678 11.56 77.00 10 Rajankunte KF790641 12.97 77.59 11 IIHR KF790663 13.13 77.48 12 Rajarajeshwarinagar KF790664 12.80 77.51 13 Kolar HQ268814 14.10 77.27 14 Kolar HQ268811 14.10 77.27 15 Banashankari KF790659 12.93 77.55 16 Chikkajala KF790661 13.16 77.63 17 Chamarajanagar KF790679 11.56 77.00 18 Kolar HQ331244 14.10 77.27 19 Bija pur KJ523177 16.83 75.71 20 Bidadi KF790640 12.80 77.40 21 Rajarajeshwarinagar KF790642 12.90 77.51 22 IIHR KF790643 13.13 77.48 23 Bidadi KF790644 12.80 77.40 24 Chikkajala KF790645 13.16 77.63 25 IIHR KF790646 13.13 77.48 26 IIHR KF790647 13.13 77.48 27 IIHR KF790648 13.13 77.48 28 Golahalli KF790649 13.13 77.59 29 IIHR KF790650 13.13 77.48 30 Dommasandra KF790651 12.88 77.75 31 Ranjantunte KF790652 12.97 77.59 32 IIHR KF790654 13.13 77.48 33 IIHR KF790655 13.13 77.48 34 Bangalore KF790666 13.04 77.59 35 Bidadi KF790656 12.80 77.40 36 Kadugedi KF790657 12.99 77.76 37 Banashankari KF790658 12.93 77.55 38 Rajajinagar KF790660 12.97 77.57 39 Anekal KF790662 12.70 77.70 40 Makali KF790665 13.09 77.39 41 Andaman KF790668 11.68 92.77 42 Gujarat KF790669 23.21 72.68 43 Kodagu KF790670 12.42 75.73 44 Maddur KF790671 12.59 77.05 45 Hunasuru KF790672 12.31 76.29 46 Mysore KF790673 12.18 76.42 47 Mysore (*Periyapatna) KF790677 12.18 76.42 48 Harangi KF790675 12.49 75.90 49 Basavanahalli KF790676 13.11 76.91 50 Channapattana KF790674 12.65 77.20 51 Chamarajanagar KF790680 11.56 77.00 52 Chamarajanagar KF790681 11.56 77.00 53 Chamarajanagar KF790682 11.56 77.00 54 Tumkur KF790683 13.20 77.08 55 Rannibennur KF790684 14.61 75.62 56 Davanagere KF790685 14.31 75.58 57 Davanagere KF790686 14.31 75.58 58 Chitradurga KF790687 14.14 76.26 59 Sirsi KF790688 14.61 74.82 60 Haveri KF790689 14.80 75.39 62 Kolar HQ268812 13.01 78.33 63 Kolar HQ331246 13.73 75.33 64 Kolar HQ331245 13.09 78.11 65 Gulbarga KJ523182 17.33 76.83 66 Raichur KJ523180 16.20 77.37 67 Chikmangaluru KJ523179 13.32 75.77 68 Dharward KJ523181 15.45 75.00 69 Belgaum KJ523178 15.85 74.50 70 Kolar (H Cross) KJ787661 13.24 77.93 71 Kolar (Malur) KJ787662 13.02 77.93 72 Kolar HQ268813 13.13 78.13 Voucher Year of SI no. Genetic group specimen collection 1 Asia-II-7 HKR01 2012 2 Asia-II-7 HKR02 2012 3 Asia-II-7 HKR03 2012 4 Asia-II-7 HKR04 2012 5 Asia-II-7 HKR05 2012 6 Asia-II-7 HKR06 2012 7 Asia-II-7 HKR20 2013 8 Asia-II-7 HKR34 2012 9 Asia-II-7 HKR45 2011 10 MEAM-1 HKR08 2012 11 MEAM-1 HKR30 2013 12 MEAM-1 HKR31 2013 13 MEAM-1 NA 2010 14 MEAM-1 NA 2010 15 Asia-II-8 HKR26 2013 16 Asia-II-8 HKR28 2013 17 Asia-II-8 HKR46 2011 18 Asia-II-8 NA 2010 19 Asia-II-8 HKR57 2013 20 Asia-1 HKR07 2012 21 Asia-1 HKR09 2012 22 Asia-1 HKR10 2012 23 Asia-1 HKR11 2012 24 Asia-1 HKR12 2012 25 Asia-1 HKR13 2012 26 Asia-1 HKR14 2012 27 Asia-1 HKR15 2012 28 Asia-1 HKR16 2013 29 Asia-1 HKR17 2013 30 Asia-1 HKR18 2013 31 Asia-1 HKR19 2013 32 Asia-1 HKR21 2013 33 Asia-1 HKR22 2013 34 Asia-1 HKR33 2013 35 Asia-1 HKR23 2013 36 Asia-1 HKR24 2013 37 Asia-1 HKR25 2013 38 Asia-1 HKR27 2013 39 Asia-1 HKR29 2013 40 Asia-1 HKR32 2013 41 Asia-1 HKR35 2012 42 Asia-1 HKR36 2011 43 Asia-1 HKR37 2011 44 Asia-1 HKR38 2011 45 Asia-1 HKR39 2011 46 Asia-1 HKR40 2011 47 Asia-1 HKR44 2011 48 Asia-1 HKR42 2011 49 Asia-1 HKR43 2011 50 Asia-1 HKR41 2011 51 Asia-1 HKR47 2011 52 Asia-1 HKR48 2011 53 Asia-1 HKR49 2011 54 Asia-1 HKR50 2011 55 Asia-1 HKR51 2011 56 Asia-1 HKR52 2011 57 Asia-1 HKR53 2011 58 Asia-1 HKR54 2011 59 Asia-1 HKR55 2011 60 Asia-1 HKR56 2011 62 Asia-1 NA 2010 63 Asia-1 NA 2010 64 Asia-1 NA 2010 65 Asia-1 HKR62 2013 66 Asia-1 HKR60 2013 67 Asia-1 HKR59 2013 68 Asia-1 HKR61 2013 69 Asia-1 HKR58 2013 70 MEAM-K HKR63 2014 71 MEAM-K HKR64 2014 72 MEAM-K NA 2010 Table 2. Maximum composite likelihood estimate of the pattern of nucleotide substitution from Bemisia tabaci populations collected on various host plants in Karnataka, India. A T C G A -- 5.24## 1.61## 14.63# T 2.98## -- 9.96# 2.34## C 2.98## 32.46# -- 2.34## G 18.62# 5.24## 1.61## -- Each entry shows the probability of substitution from one base (row) to another base (column) instantaneously. Only entries within a row should be compared. Rates of different transitional substitutions are shown in bold and those of transversional substitutions are shown in italics. Note: Rates of different transitional substitutions shown are indicated with # and those of transversional substitutions are shown in ##. Table 3. Estimates of net evolutionary divergence between groups of sequences showing pair-wise genetic distances for Bemisia tabaci populations collected on various host plants in Karnataka, India. Asia-II-7 Asia-I 0.139 MEAM-1 0.173 0.161 Asia-II-8 0.119 0.152 0.161 MEAM-l-Karnataka 0.153 0.164 0.074 0.157 Asia-III 0.135 0.078 0.174 0.163 0.175 Asia-II-10 0.109 0.137 0.172 0.120 0.155 Asia-II-1 0.092 0.157 0.172 0.133 0.160 Asia-II-2 0.111 0.144 0.129 0.120 0.130 Asia-II-3 0.122 0.176 0.174 0.118 0.165 Asia-II-4 0.129 0.169 0.160 0.142 0.166 Asia-II-5 0.109 0.173 0.172 0.117 0.095 Asia-II-6 0.096 0.159 0.171 0.136 0.124 Asia-II-9 0.108 0.169 0.169 0.118 0.155 Australia 0.156 0.117 0.163 0.155 0.153 Australia-Indonesia 0.178 0.146 0.174 0.176 0.166 China-I 0.145 0.128 0.161 0.148 0.168 Indian_Ocean 0.165 0.152 0.076 0.169 0.126 Italy 0.124 0.145 0.153 0.128 0.155 Mediterranean 0.170 0.171 0.052 0.164 0.110 New-World-1 0.159 0.134 0.139 0.163 0.166 Subgroup-Af-1 0.189 0.196 0.164 0.184 0.195 Subgroup-Af-2 0.205 0.186 0.165 0.187 0.198 Subgroup-Af-3 0.199 0.172 0.172 0.176 0.197 Subgroup-Af-4 0.195 0.183 0.175 0.189 0.201 Uganda 0.229 0.228 0.202 0.207 0.209 MEAM-2 0.162 0.160 0.041 0.150 0.098 China-II 0.135 0.127 0.153 0.154 0.144 China-III 0.151 0.150 0.156 0.135 0.154 Asia-II-7 Asia-I MEAM-1 Asia-II-8 MEAM-l-Karnataka Asia-III Asia-II-10 0.139 Asia-II-1 0.158 0.131 Asia-II-2 0.145 0.108 0.067 Asia-II-3 0.166 0.107 0.133 0.075 Asia-II-4 0.158 0.119 0.125 0.051 0.048 Asia-II-5 0.168 0.139 0.106 0.107 0.136 Asia-II-6 0.147 0.116 0.107 0.118 0.136 Asia-II-9 0.160 0.098 0.121 0.091 0.050 Australia 0.124 0.154 0.148 0.130 0.152 Australia-Indonesia 0.139 0.170 0.179 0.151 0.176 China-I 0.122 0.134 0.156 0.135 0.156 Indian_Ocean 0.171 0.168 0.167 0.153 0.176 Italy 0.131 0.131 0.140 0.134 0.134 Mediterranean 0.185 0.177 0.179 0.150 0.182 New-World-1 0.152 0.147 0.165 0.148 0.157 Subgroup-Af-1 0.187 0.180 0.193 0.182 0.184 Subgroup-Af-2 0.191 0.184 0.205 0.183 0.175 Subgroup-Af-3 0.180 0.178 0.190 0.180 0.188 Subgroup-Af-4 0.185 0.172 0.202 0.188 0.193 Uganda 0.220 0.198 0.212 0.211 0.237 MEAM-2 0.166 0.167 0.163 0.137 0.169 China-II 0.130 0.134 0.157 0.139 0.158 China-III 0.142 0.138 0.161 0.140 0.162 Asia-II-7 Asia-I MEAM-1 Asia-II-8 MEAM-l-Karnataka Asia-III Asia-II-10 Asia-II-1 Asia-II-2 Asia-II-3 Asia-II-4 Asia-II-5 0.133 Asia-II-6 0.141 0.067 Asia-II-9 0.080 0.120 0.126 Australia 0.142 0.133 0.150 0.147 Australia-Indonesia 0.180 0.172 0.174 0.163 0.136 China-I 0.154 0.147 0.141 0.148 0.147 Indian_Ocean 0.183 0.177 0.175 0.163 0.180 Italy 0.148 0.142 0.129 0.123 0.131 Mediterranean 0.186 0.187 0.171 0.165 0.191 New-World-1 0.159 0.162 0.173 0.145 0.164 Subgroup-Af-1 0.198 0.202 0.199 0.178 0.182 Subgroup-Af-2 0.191 0.205 0.190 0.164 0.175 Subgroup-Af-3 0.197 0.216 0.204 0.180 0.189 Subgroup-Af-4 0.203 0.208 0.206 0.187 0.183 Uganda 0.239 0.218 0.225 0.212 0.233 MEAM-2 0.171 0.172 0.157 0.162 0.174 China-II 0.152 0.142 0.145 0.144 0.146 China-III 0.168 0.165 0.160 0.140 0.150 Asia-II-7 Asia-I MEAM-1 Asia-II-8 MEAM-l-Karnataka Asia-III Asia-II-10 Asia-II-1 Asia-II-2 Asia-II-3 Asia-II-4 Asia-II-5 Asia-II-6 Asia-II-9 Australia Australia-Indonesia China-I 0.138 Indian_Ocean 0.180 0.165 Italy 0.122 0.123 0.153 Mediterranean 0.180 0.168 0.070 0.161 New-World-1 0.179 0.150 0.156 0.128 0.148 Subgroup-Af-1 0.180 0.186 0.187 0.155 0.181 Subgroup-Af-2 0.186 0.198 0.176 0.149 0.176 Subgroup-Af-3 0.183 0.189 0.183 0.165 0.185 Subgroup-Af-4 0.181 0.199 0.182 0.156 0.180 Uganda 0.207 0.206 0.213 0.209 0.207 MEAM-2 0.177 0.154 0.069 0.148 0.040 China-II 0.142 0.044 0.160 0.121 0.173 China-III 0.115 0.123 0.191 0.142 0.179 Asia-II-7 Asia-I MEAM-1 Asia-II-8 MEAM-l-Karnataka Asia-III Asia-II-10 Asia-II-1 Asia-II-2 Asia-II-3 Asia-II-4 Asia-II-5 Asia-II-6 Asia-II-9 Australia Australia-Indonesia China-I Indian_Ocean Italy Mediterranean New-World-1 Subgroup-Af-1 0.149 Subgroup-Af-2 0.147 0.076 Subgroup-Af-3 0.151 0.072 0.061 Subgroup-Af-4 0.150 0.078 0.072 0.071 Uganda 0.194 0.190 0.196 0.194 MEAM-2 0.144 0.167 0.163 0.175 China-II 0.144 0.190 0.195 0.189 China-III 0.174 0.176 0.195 0.191 Asia-II-7 Asia-I MEAM-1 Asia-II-8 MEAM-l-Karnataka Asia-III Asia-II-10 Asia-II-1 Asia-II-2 Asia-II-3 Asia-II-4 Asia-II-5 Asia-II-6 Asia-II-9 Australia Australia-Indonesia China-I Indian_Ocean Italy Mediterranean New-World-1 Subgroup-Af-1 Subgroup-Af-2 Subgroup-Af-3 Subgroup-Af-4 Uganda 0.202 MEAM-2 0.170 0.195 China-II 0.204 0.209 0.159 China-III 0.196 0.205 0.157 0.120 Analyses were conducted using the maximum composite likelihood model (Tajima & Nei 1984). The analysis involved 135 nucleotide sequences. All positions containing gaps and missing data were eliminated. There were 620 positions in the final dataset. Evolutionary analyses were conducted in MEGA5 (Tamura et al. 2011).
Please note: Some tables or figures were omitted from this article.
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|Title Annotation:||Research Papers|
|Author:||Roopa, H.K.; Asokan, R.; Rebijith, K.B.; Hande, Ranjitha H.; Mahmood, Riaz; Kumar, N.K. Krishna|
|Date:||Dec 1, 2015|
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