Development of abamectin resistance in Tetranychus urticae in Australian cotton and the establishment of discriminating doses for T. lambi.

Since the 1980s Tetranychus urticae Koch has dominated Australian cotton due to its ability to develop resistance. Here we give screening data for a range of chemicals tested against T. urticae including abamectin, bifenthrin, diafenthiuron, etoxazole and propargite and speculate why abamectin resistance emerged without warning. Abamectin resistance was not detected in T. urticae in Australian cotton before season 2007-2008 when a few resistant individuals were detected in a single strain. Resistance was detected again in season 2010-2011 and continued to be detected in every subsequent season comprising 80% of strains tested in 2018-2019. We speculate the reason may relate to prophylactic abamectin use to prevent mite flare with Creontiades dilutes Stål mirid sprays. With the introduction of transgenic Bt-cotton, spraying significantly reduced and anecdotally Tetranychus lambi became more abundant. Although T. lambi may now be more common than T. urticae its response to chemical controls is completely unknown. Tetranychus lambi conspecific dose responses were established to support resistance monitoring against abamectin, diafenthiuron and propargite that generated discriminating dose (DD) estimates of 0.0007 g/L abamectin, 0.03 g/L diafenthiuron and 0.7 g/L propargite. These DD were used in season 2018-2019 but resistance was not detected against any product including abamectin. The reason why T. lambi may now dominate despite T. urticae being still resistant is speculated and thought related to the progressive reduction in insecticide use in Australian cotton and/or the changing weed complex in the transgenic cotton era.

Multiple invasions of a generalist herbivore-Secondary contact between two divergent lineages of Nezara viridula Linnaeus in Australia.

The presence of distinct evolutionary lineages within herbivorous pest insect taxa requires close attention. Scientific understanding, biosecurity planning and practice, and pest management decision-making each suffer when such situations remain poorly understood. The pest bug Nezara viridula Linnaeus has been recorded from numerous host plants and has two globally distributed mitochondrial (mtDNA) lineages. These mtDNA lineages co-occur in few locations globally, and the consequences of their divergence and recent secondary contact have not been assessed. We present evidence that both mtDNA lineages of N. viridula are present in Australia and their haplotype groups have a mostly separate distribution from one another. The north-western population has only Asian mtDNA haplotypes, and the population with an eastern distribution is characterized mostly by European mtDNA haplotypes. Haplotypes of both lineages were detected together at only one site in the north of eastern Australia, and microsatellite data indicate that this secondary contact has resulted in mating across the lineages. Admixture and the movement of mtDNA haplotypes outside of this limited area of overlap has not, however, been extensive. Some degree of mating incompatibility or differences in the climatic requirements and tolerances of the two lineages, and perhaps a combination of these influences, might limit introgression and the movement of individuals, but this needs to be tested. This work provides the foundation for further ecological investigation of the lineages of N. viridula, particularly the consequences of admixture on the ecology of this widespread pest. We propose that for now, the Asian and European lineages of N. viridula would best be investigated as subspecies, so that "pure" and admixed populations of this bug can each be considered directly with respect to management and research priorities.

The Management of Insect Pests in Australian Cotton: An Evolving Story.

The Australian cotton industry progressively embraced integrated pest management (IPM) to alleviate escalating insecticide resistance issues. A systems IPM approach was used with core principles that were built around pest ecology/biology and insecticide resistance management; together, these were integrated into a flexible, year-round approach that facilitated easy incorporation of new science, strategies, and pests. The approach emphasized both strategic and tactical elements to reduce pest abundance and rationalize decisions about pest control, with insecticides as a last resort. Industry involvement in developing the approach was vital to embedding IPM within the farming system. Adoption of IPM was facilitated by the introduction of Bt cotton, availability of selective insecticides, economic validation, and an industry-wide extension campaign. Surveys indicate IPM is now embedded in industry, confirming the effectiveness of an industry-led, backed-by-science approach. The amount of insecticide active ingredient applied per hectare against pests has also declined dramatically. Though challenges remain, pest management has transitioned from reactively attempting to eradicate pests from fields to proactively managing them year-round, considering the farm within the wider landscape.

Effects of single and dual species herbivory on the behavioral responses of three thrips species to cotton seedlings.

This study investigated the olfactory responses of 3 thrips species [Frankliniella schultzei Trybom, F. occidentalis Pergrande and Thrips tabaci Lindeman (Thysanoptera: Thripidae)] to cotton seedlings [Gossypium hirsutum L. (Malvales: Malvaceae)] simultaneously damaged by different combinations of herbivores. Cotton seedlings were damaged by foliar feeding Tetranychus urticae Koch (Trombidiforms: Tetranychidae), Helicoverpa armigera Hübner (Lepidoptera: Noctuidae), Aphis gossypii Glover (Hemiptera: Aphididae) or root feeding Tenebrio molitor L. (Coleoptera: Tenebrionidae). Thrips responses to plants simultaneously damaged by 2 species of herbivore were additive and equivalent to the sum of the responses of thrips to plants damaged by single herbivore species feeding alone. For example, F. occidentalis was attracted to T. urticae damaged plants but more attracted to undamaged plants than to plants damaged by H. armigera. Plants simultaneously damaged by low densities of T. urticae and H. armigera repelled F. occidentalis but as T. urticae density increased relative to H. armigera density, F. occidentalis attraction to coinfested plants increased proportionally. Thrips tabaci did not discriminate between undamaged plants and plants damaged by H. armigera but were attracted to plants damaged by T. urticae alone or simultaneously damaged by T. urticae and H. armigera. Olfactometer assays showed that simultaneous feeding by 2 herbivores on a plant can affect predator-prey interactions. Attraction of F. occidentalis to plants damaged by its T. urticae prey was reduced when the plant was simultaneously damaged by H. armigera, T. molitor, or A. gossypii and F. schultzei was more attracted to plants simultaneously damaged by T. urticae and H. armigera than to plants damaged by T. urticae alone. We conclude that plant responses to feeding by 1 species of herbivore are affected by responses to feeding by other herbivores. These plant-mediated interactions between herbivore complexes affect the behavioral responses of thrips which vary between species and are highly context dependent.

Enhancing Integrated Pest Management in GM Cotton Systems Using Host Plant Resistance.

Cotton has lost many ancestral defensive traits against key invertebrate pests. This is suggested by the levels of resistance to some pests found in wild cotton genotypes as well as in cultivated landraces and is a result of domestication and a long history of targeted breeding for yield and fiber quality, along with the capacity to control pests with pesticides. Genetic modification (GM) allowed integration of toxins from a bacteria into cotton to control key Lepidopteran pests. Since the mid-1990s, use of GM cotton cultivars has greatly reduced the amount of pesticides used in many cotton systems. However, pests not controlled by the GM traits have usually emerged as problems, especially the sucking bug complex. Control of this complex with pesticides often causes a reduction in beneficial invertebrate populations, allowing other secondary pests to increase rapidly and require control. Control of both sucking bug complex and secondary pests is problematic due to the cost of pesticides and/or high risk of selecting for pesticide resistance. Deployment of host plant resistance (HPR) provides an opportunity to manage these issues in GM cotton systems. Cotton cultivars resistant to the sucking bug complex and/or secondary pests would require fewer pesticide applications, reducing costs and risks to beneficial invertebrate populations and pesticide resistance. Incorporation of HPR traits into elite cotton cultivars with high yield and fiber quality offers the potential to further reduce pesticide use and increase the durability of pest management in GM cotton systems. We review the challenges that the identification and use of HPR against invertebrate pests brings to cotton breeding. We explore sources of resistance to the sucking bug complex and secondary pests, the mechanisms that control them and the approaches to incorporate these defense traits to commercial cultivars.

Effect of the postfeeding interval on olfactory responses of thrips to herbivore-induced cotton plants.

We investigated the responses of 3 thrips species, Frankliniella schultzei Trybom, F. occidentalis Pergrande, and Thrips tabaci Lindeman (Thysanoptera: Thripidae) to herbivore-damaged and undamaged cotton seedlings (Gossypium hirsutum L. [Malvales: Malvaceae]) at a range of time intervals following damage by adult Tetranychus urticae (Koch), adult T. ludeni (Zacher) (Acari: Tetranychidae) or Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae in olfactometer assays. The intensity/frequency of the response of thrips to herbivore-induced plants decreased with time and ultimately disappeared in all cases; however, the rate at which the response declined was related to the herbivore species that inflicted the damage. All 3 species of thrips were attracted to plants damaged by T. urticae for longer than they were to plants damaged by T. ludeni. The duration for which damaged plants remained attractive was also affected by the degree of damage inflicted on cotton seedlings. For example, F. schultzei was attracted to plants damaged by a higher density of two-spotted spider mites (100/plant) for much longer than to plants damaged by a lower density of these mites (50/plant). The results reinforce previous studies that demonstrate that arrangement of variables influences the responses of thrips to their herbivore-induced cotton host plants. Results also show that these responses are variable in time following herbivore damage to cotton plants, which further demonstrates how difficult it is to generalize about the functional significance of these interactions.

Effects of Season and Management of Irrigated Cotton Fields on Collembola (Hexapoda) in New South Wales, Australia.

The effects of production practices on the relative abundance of springtails (Collembola) in irrigated cotton fields of northern New South Wales (NSW) were studied over 2 yr to examine effects of farm management on these decomposer organisms. Pitfall trapping and soil core extraction was undertaken in both pseudoreplicated plots within whole fields on cotton farms and on experimental replicate plots of Envirofeast cotton and Lucerne. The relative abundance of surface-active springtails in cotton rows and densities of soil species from the rhizosphere were calculated. Twenty-three species of Collembola were collected from 5 fields, 19 in pitfall traps, and 11 in soil cores. Five species, Setogaster sp., Proisotoma minuta, Entomobrya unostrigata, Entomobrya multifasciata grp, and Lepidobrya sp. were numerically dominant on the ground at 86-96% of individuals and Mesaphorura sp., Folsomides parvulus, and Hemisotoma thermophila grp dominant in the soil. Native grassland samples contained 15 species of which a probable 10 were native and 8 were not found in cotton. Nineteen species of the 24 species identified from cotton were predominantly fungal feeders. Highest catches of Collembola occurred after flowering and soil Collembola increased with depth and during cotton growth on unsprayed plots but decreased on sprayed plots. Surface soil moistures affected daily catch rates with decomposing residues, crop stage, predator abundance, and season as secondary factors. Insecticide (endosulfan, pyrethroid, carbamate, and organophosphate) and predator effects were either negligible or unclear depending on the factor involved. Springtails appear to be predominately food limited during times of adequate soil moisture in cotton fields.

Lack of adaptation to a new host in a generalist herbivore: implications for host plant resistance to twospotted spider mites in cotton.

BACKGROUND: The twospotted spider mite (Tetranychus urticae Koch) is an important pest of cotton. This pest has a broad host range, but when changing between hosts an initial decline in fitness often occurs. This is usually followed by an increase in fitness after rapid adaptation to the new host, usually within five generations. RESULTS: The generality of this adaptive response was tested by assessing elements of fitness when mites were reared on a host to which they were adapted (Gossypium hirsutum L. cv. Sicot 71) or on a new host, Gossypium arboreum L. (accession BM13H). In a first experiment, mites reared on the new host for ten generations showed declining immature survival compared with those reared on the adapted host. In a second experiment, the intrinsic capacity for increase of mites cultured on the new host for six generations was significantly lower than that of mites cultured on the adapted host for six generations and then transferred to the new host. Hence, exposure to the new host for six or ten generations resulted in declining fitness. CONCLUSION: This 'negative adaptation' indicates robust antibiosis traits in G. arboreum accession BM13H, which therefore have value in developing mite-resistant G. hirsutum cultivars.

Jasmonic acid is associated with resistance to twospotted spider mites in diploid cotton (Gossypium arboreum).

The twospotted spider mite (Tetranychus urticae Koch) is capable of dramatically reducing the yield of cotton crops and is often difficult and expensive to control. This study investigated and compared two important plant hormones, jasmonic acid (JA) and salicylic acid (SA), as constitutive and/or induced defence response components in a mite susceptible commercial cotton cultivar, Sicot 71 (Gossypium hirsutum L.) and a resistant diploid cotton BM13H (Gossypium arboreum L.). Foliar application of JA and methyl jasmonate (MeJA) reduced the mite population and leaf damage but application of other potential elicitors, SA and methyl salicylate (MeSA) did not. The concentrations of JA and SA in leaf tissues of induced and non-induced Sicot 71 and BM13H were quantified by liquid chromatography coupled to electrospray ionisation tandem mass spectrometry (LC-ESI-MS/MS). The JA content was constitutively higher in BM13H than Sicot 71 and also highly induced by mite infestation in BM13H but not in Sicot 71. However, SA was not significantly induced in either BM13H or Sicot 71. The expression levels of JA related genes, LOX, AOS and OPR were measured by quantitative PCR and elevated expression levels of JA related genes were detected in mite-infested BM13H. Therefore, JA and MeJA were implicated as key biochemical components in both the constitutive and induced defence responses of BM13H to spider mites.

How predictable are the behavioral responses of insects to herbivore induced changes in plants? Responses of two congeneric thrips to induced cotton plants.

Changes in plants following insect attack are referred to as induced responses. These responses are widely viewed as a form of defence against further insect attack. In the current study we explore whether it is possible to make generalizations about induced plant responses given the unpredictability and variability observed in insect-plant interactions. Experiments were conducted to test for consistency in the responses of two congeneric thrips, Frankliniella schultzei Trybom and Frankliniella occidentalis Pergrande (Thysanoptera: Thripidae) to cotton seedlings (Gossypium hirsutum Linneaus (Malvales: Malvaceae)) damaged by various insect herbivores. In dual-choice experiments that compared intact and damaged cotton seedlings, F. schultzei was attracted to seedlings damaged by Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), Tetranychus urticae (Koch) (Trombidiforms: Tetranychidae), Tenebrio molitor Linnaeus (Coleoptera: Tenebrionidae), F. schultzei and F. occidentalis but not to mechanically damaged seedlings. In similar tests, F. occidentalis was attracted to undamaged cotton seedlings when simultaneously exposed to seedlings damaged by H. armigera, T. molitor or F. occidentalis. However, when exposed to F. schultzei or T. urticae damaged plants, F. occidentalis was more attracted towards damaged plants. A quantitative relationship was also apparent, F. schultzei showed increased attraction to damaged seedlings as the density of T. urticae or F. schultzei increased. In contrast, although F. occidentalis demonstrated increased attraction to plants damaged by higher densities of T. urticae, there was a negative relationship between attraction and the density of damaging conspecifics. Both species showed greater attraction to T. urticae damaged seedlings than to seedlings damaged by conspecifics. Results demonstrate that the responses of both species of thrips were context dependent, making generalizations difficult to formulate.

Fitness of twospotted spider mites is more affected by constitutive than induced resistance traits in cotton (Gossypium spp.).

BACKGROUND: Life history parameters are useful tools for comparing the fitness of pests on different host plants. This study compared life history parameters of twospotted spider mites (Tetranychus urticae Koch) on two resistant cotton Gossypium genotypes (BM13H and Sipima 280) and one susceptible genotype (Sicot 71). The effects of both constitutive and induced defences were assessed. RESULTS: Mites reared on the resistant genotypes had longer immature development times, lower immature survival and reduced adult fecundity. Mites reared on BM13H that had been induced by prior exposure to mites had a small additional decrease in adult fecundity. The contribution to mite resistance of constitutive resistance mechanisms was much greater than induced responses. The effect of morphological constitutive defences was minor, implicating biochemical defences as the major mite-resistance mechanism. Sensitivity analysis and a population development study using life history parameters of mites showed that a lower immature survival rate on resistant genotypes had the greatest effect on mite fitness and population development. CONCLUSION: Use of life history parameters provided valuable insight into the mite-resistance mechanisms of these Gossypium genotypes. Further, the results largely explained mite population development on these genotypes in the field.

Evidence of superclones in Australian cotton aphid Aphis gossypii Glover (Aphididae: Hemiptera).

BACKGROUND: Aphis gossypii is an important pest of cotton that has developed resistance to many chemicals used for its control. Any lack of understanding of its genetic structure, resistance status and host plant specialisation hampers effective management. RSULTS: Eight microsatellite markers were genotyped for a collection of Australian A. gossypii field isolates from 55 plant species from major Australian cotton-producing regions. The aphid's pirimicarb resistance status linked to the ACE1 (acetylcholinesterase) S431F mutation was determined by PCR-RFLP. Overall, the genetic diversity was low and there were only 13 multilocus genotype (MLG) groups found in a total of 936 aphids, suggesting asexual reproduction. Three MLGs (Aust-01, Aust-02 and Aust-04) represented 78% of all aphids tested. MLGs Aust-01 (41%) and Aust-02 (18%) were linked to the ACE1 S431F mutation and found on cotton and a range of hosts. Aust-04 (19%) hosted mainly on cotton (but also Asteraceae and Malvaceae) was predominantly susceptible to pirimicarb. Given their abundance and widespread occurrence, these three clones were considered to be superclones. CONCLUSION: The study demonstrated that any strategy to control A. gossypii and manage pirimicarb resistance should target A. gossypii strains of all MLG types residing on any plant species and not just cotton

Contradictions in host plant resistance to pests: spider mite (Tetranychus urticae Koch) behaviour undermines the potential resistance of smooth-leaved cotton (Gossypium hirsutum L.).

BACKGROUND: Two-spotted spider mites (Tetranychus urticae Koch) oviposit near leaf veins or in leaf folds on the undersides of cotton (Gossypium hirsutum L.) leaves where the humid boundary layer offers protection from desiccation. The authors predicted that the boundary layer of glabrous cotton leaves should be shallower than that of hairy leaves, providing some resistance to mites. The dynamics of mite populations, leaf damage, leaf gas exchange and crop yield on two leaf hair isolines (smooth versus hairy) in two genetic backgrounds was assessed. RESULTS: Mite colonies developed faster on the hairy leaf isolines, but leaf damage per mite was higher in smooth leaf isolines, indicating more intense damage. A 50% reduction in photosynthesis on the hairy isolines required 1.8 times more mites than smooth leaves. The yield of cotton was reduced in + mite treatments, but the magnitude of reduction was similar for hairy and smooth isolines. CONCLUSION: Paradoxically, the relative inhospitality of glabrous leaves may have induced mites to concentrate in protected leaf sections, causing more localised and more severe damage, negating the yield benefits from fewer mites. These results highlight interactions between leaf microenvironment, pest behaviour and plant productivity that may have implications for other instances of plant resistance.

Recovery of leaf area through accelerated shoot ontogeny in thrips-damaged cotton seedlings.

BACKGROUND AND AIMS: Leaf area of cotton seedlings (Gossypium hirsutum) can be reduced by as much as 50 % by early season thrips infestations, but it is well documented that plants can regain the difference in leaf area once infestation ceases. The processes involved in the recovery have not been identified. Hypotheses include enhancement of the photosynthetic rate of the damaged leaves, more efficient leaf construction (i.e. more leaf area per unit of dry matter invested in new leaves), and more branching. METHODS: This 2-year field study examined these hypotheses and found that thrips-affected plants recovered from a 30 % reduction in total leaf area. During the recovery period, repeated measurements of gas exchange, leaf morphology and individual leaf areas at all nodes were made to assess their contribution to the recovery. KEY RESULTS: Recovery was not achieved through the previously proposed mechanisms. The pattern of nodal development indicated that the duration of leaf expansion of the smaller deformed leaves was shorter than that of control leaves, possibly because they had fewer cells. The production and expansion of healthy upper node leaves in thrips-affected plants could, therefore, begin sooner, about 1-2.5 nodes in advance of control plants. The proposed process of recovery was evident but weaker in the second year where thrips numbers were higher. CONCLUSIONS: It is concluded that thrips-affected plants overcame the leaf area disparity through an accelerated ontogeny of main stem leaves. By completing the expansion of smaller but normally functioning lower node leaves earlier, resources were made available to the unfolding of larger upper node leaves in advance of control plants. The generality of this mode of plant resistance in pest damage remains to be determined.