Status of two cryptic species, Typhlodromus exhilaratus Ragusa and Typhlodromus phialatus Athias-Henriot (Acari: Phytoseiidae): consequences for taxonomy

Typhlodromus phialatus and T. exhilaratus are morphologically close species. Their differentiation is based on the shape of the insemination apparatus and on idiosomal setae length. However, the setae length values are often intermediate between these two species and do not allow accurate identification. Furthermore, the handful of differences in insemination apparatus shape are also questionable as a means of differentiation. Synonymy between these species has thus been questioned. Three experiments were carried out. Idiosomal seta length measurements, molecular tests and cross experiments were conducted for three populations, identified as T. exhilaratus and T. phialatus according to the shape of their insemination apparatus. The results show that the variation range of seta lengths is great and that these criteria do not allow accurate separation of these populations into two species. However, molecular tests show a species-level nucleotide differentiation between them. Cross experiments confirm this result, showing complete reproduction incompatibility between the mites bearing different insemination apparatus shapes. Therefore, T. exhilaratus and T. phialatus could be considered to be two valid species and the insemination apparatus could be considered as a pertinent diagnostic criterion at the specific level.     

Differential impact of egg predation by Zetzellia mali (Acari: Stigmaeidae) on Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

The differential impact of Zetzellia mali on the phytoseiids Metaseiulus occidentalis and Typhlodromus pyri was studied in the laboratory and by analysis of population from orchard plots that contained either phytoseiid, similar numbers of prey mites and high or low densities of Z. mali. Five hypotheses were evaluated to explain why Z. mali had more impact on M. occidentalis in the field than on T. pyri. Given equal opportunity, Z. mali adult females did not consume more M. occidentalis eggs than T. pyri eggs nor did adult females of either phytoseiid inflict greater mortality on Z. mali eggs or larvae through attack or consumption. There was no difference in the within-tree association of Z. mali adult females with eggs of either phytoseiid species nor were there differences in the way prey mites (all life stages) were spatially partitioned between adult female Z. mali as compared with adults and deutonymphs (combined) of either phytoseiid. The foraging area of adult female Z. mali and the oviposition locations of the two phytoseiids from both field and laboratory data were compared using spatial statistics. Metaseiulus occidentalis laid significantly more eggs in the primary foraging area of adult female Z. mali than T. pyri. This was the only factor identified which may explain the greater impact of Z. mali on M. occidentalis. The impact of these interspecific effects on the persistence of predatory mite guilds and biological control are discussed.     

Interspecific competition and predation between immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Interspecific competition and predation in immature Amblyseius fallacis (Garman), Amblyseius andersoni Chant, Typhlodromus occidentalis (Nesbitt) and Typhlodromus pyri Scheuten were examined in small cages at three egg densities (0, 20 and 80) of two-spotted spider mite, Tetranychus urticae Koch, in the laboratory at 25±1°C,80% RH and 16L: 8D photoperiod. For the six possible between-species comparisons, the large polyphagous A. andersoni always outcompeted the other three predator species, which were either smaller and/or less polyphagous; the small oligophagous T. occidentalis was always eliminated by the other three predator species, which were either larger and/or more polyphagous. The small and polyphagous T. pyri tied with the large and oligophagous A. fallacis. The outcome of the interaction was generally similar at the three prey densities except in (1) the A. fallacis-A. andersoni system where the advantage of A. andersoni over A. fallacis was reduced when 20 or 80 eggs per cage were present at the start of the interaction and (2) the A. fallacis-T. occidentalis system where the advantage of A. fallacis over T. occidentalis increased with prey density. This study indicates that predator size, predator degree of polyphagy and prey density can affect the competitiveness of immature phytoseiids.     

Effects of chlorpyrifos and sulfur on spider mites (Acari: Tetranychidae) and their natural enemies

In many agricultural systems spider mites are believed to be induced pests, only reaching damaging densities after pesticides decimate predator populations. Wine grapes typically receive two types of pesticides, insecticides and fungicides. Chemicals in either class could impact spider mite densities both directly through spider mite mortality, and indirectly by negatively affecting natural enemies. The impact of a broad-spectrum insecticide (chlorpyrifos) and an inorganic fungicide (sulfur) on mites and their natural enemies was monitored in replicate open-field experiments conducted in an abandoned vineyard in Washington State. In both experiments, chemicals were applied within a 2 × 2 factorial design, allowing assessment of both main and interactive effects of the two chemicals. Following typical management practices on wine grapes in Washington State, we made a single insecticide application early in the season, but repeatedly applied sulfur throughout the season. In the absence of sulfur, chlorpyrifos application led to higher spider mite densities. The main effect of chlorpyrifos appeared to be indirect, perhaps mediated through mortality of generalist phytoseiid mites; generalists appeared to be unable to recover following even a single insecticide application, while there was no evidence for harmful effects of chlorpyrifos on specialist phytoseiid mites. Sulfur had direct suppressive effects on both pest and predatory mites, although in the second experiment the suppressive effect of sulfur on spider mites was weaker when chlorpyrifos was also applied. These field experiments suggest that a complex mix of direct and indirect effects of the two chemicals impacted spider mite population dynamics in our system.     

Predation Preference of Typhlodromus Pyri and Kampimodromus Aberrans (Acari: Phytoseiidae) When Offered Con- and Heterospecific Immature Life Stages

The predation preference of singly caged adult females and nymphs of Typhlodromus pyri and Kampimodromus aberrans for con- or heterospecific immature stages as prey was tested in the laboratory. Both polyphagous predatory mite species have been previously shown to interact directly through predation on each other. The present study demonstrated that the adult females of T. pyri and K. aberrans are able to discriminate between con- and heterospecific larvae and protonymphs and that they prefer to prey upon heterospecifics when given the choice. Hunger did not reduce the propensity of the females to prefer heterospecifics over conspecifics. For proto-and deutonymphs the trends followed the results obtained with the adult females, but the preference for heterospecifics was not distinct enough to be significant. In competitive situations reciprocal predation may be a crucial mechanism in the interaction of polyphagous phytoseiid species and may contribute significantly to population persistence. The results are discussed with regard to possible associations between the ability to discriminate con- and heterospecifics and the type of feeding specialization (generalists versus specialists).     

Comparative Behavioural Studies of Larval and Adult Stages of the Phytoseiids (Acari: Mesostigmata) Typhlodromus Athiasae and Neoseiulus Californicus

We compared the behaviours of the indigenous Typhlodromus athiasae Porath and Swirski and the exotic Neoseiulus californicus (McGregor) (= Amblyseius chilenensis Dosse) relative to their persistence in apple orchards in Israel. We studied (1) larval feeding, walking, intraspecific interactions (cannibalism, touch-avoidance responses and/or touching with palps and tarsi) and tendency to aggregate (when resting), (2) predation and cannibalism on phytoseiid eggs by young females and (3) the effects of starvation for 10 days on young females relative to ambulation speed, longevity, fecundity, progeny survival and sex ratio. Larvae of T. athiasae were almost inert, did not feed and hardly walked or interacted whereas larvae of N. californicus fed, walked and interacted, mainly by touching with palps and tarsi. No cannibalism in the larval stage was observed for either species. The presence of prey increased the larval walking and intraspecific interactions of N. californicus but not of T. athiasae. Egg predation by adult females of both species was substantially higher than cannibalism, implying that both are capable of distinguishing their eggs from those of other species. Soaking eggs for 30 min in deionized water increased cannibalism in both species. During the 10 days of starvation, the ambulation speed of adult female N. californicus ranged from 1.8 to 10.1 times that of T. athiasae. The 50% lethal time value (LT₅₀) of T. athiasae (6.0 days) was significantly lower than that of N. californicus (10.4 days). None of the starved T. athiasae recuperated following the reintroduction of prey, whereas 75% of N. californicus did and oviposited after 2 days. These traits should enable N. californicus to persist when prey is scarce; however the selective predation of N. californicus eggs by T. athiasae could prevent establishment of N. californicus. The degree of specialization of these two predators is discussed.     

Olfactory response of Typhlodromus pyri (Acari: Phytoseiidae) to synthetic methyl salicylate in laboratory bioassays

The herbivore‐induced plant volatile (HIPV) methyl salicylate (MeSA) is widely present in the chemical profile of several plant species and is known to attract natural enemies, including predatory mites. In this study, the response of Typhlodromus pyri, a key predator of pest mites in west coast vineyards, to synthetically produced MeSA was tested using a Y‐tube olfactometer in laboratory bioassays. Six doses ranging from 0.002 to 200 μg of MeSA diluted in 0.1 ml hexane were tested. Significantly higher proportions of T. pyri preferred MeSA at doses 0.02, 0.2 and 20 μg. No differences in response to MeSA were detected at the highest (200 μg), intermediate (2 μg) and lowest (0.002 μg) doses. Mite response to MeSA was a function of dose when fitting polynomial and logistic regression models using dose and square of the log dose prediction factors. Results indicate that synthetic MeSA may be applied to attract predatory arthropod populations in vineyards to enhance biological control of pest mites.     

有益真绥螨与巴氏新小绥螨的集团内捕食和同类相残作用

【目的】有益真绥螨Euseius utilis是北方地区广泛分布的一种多食性植绥螨,而巴氏新小绥螨Neoseiulus barkeri目前在我国广泛应用于农业害虫的生物防治中。本研究旨在对巴氏新小绥螨在本地的应用进行风险评估及为与有益真绥螨的联合释放提供依据。【方法】本研究在室内通过一系列实验,比较了实验室饲养的巴氏新小绥螨和采自内蒙古农业大学校园的有益真绥螨的雌成螨对同种或异种未成熟螨的捕食量、存活时间及产卵量的差异,检测了有益真绥螨与巴氏新小绥螨两种植绥螨相互之间的攻击强度以及种内和种间的相互作用。【结果】两种植绥螨都难以刺吸同种或异种植绥螨的卵,而对同种或异种植绥螨幼螨的捕食量最大,其次是对若螨。在无共同食物的情况下,有益真绥螨雌成螨对同种植绥螨幼螨的攻击性比对异种植绥螨幼螨的攻击性强(BreslowDay检验:χ~2=13.84,df=1,P0.001),且有益真绥螨对同种植绥螨幼螨的捕食量(9.10±1.65头)高于对异种植绥螨幼螨的捕食量(5.31±1.43头)(T检验:t=5.487,P0.001),巴氏新小绥螨对异种植绥螨幼螨的捕食量(7.48±0.75头)高于对同种植绥螨幼螨的捕食量(4.75±0.58头)(T检验:t=9.110,P0.05)。【结论】有益真绥螨更倾向于捕食同种幼螨而发生同类相残;巴氏新小绥螨更倾向于捕食异种幼螨而发生集团内捕食。     

Juvenile Survival and Development in Euseius Finlandicus, Typhlodromus Pyri and Kampimodromus Aberrans (Acari: Phytoseiidae) Feeding on Con- and Heterospecific Immatures

Juvenile survival and development in Euseius finlandicus (Oudemans), Typhlodromus pyri Scheuten and Kampimodromus aberrans (Oudemans) feeding on con- and heterospecific phytoseiid immatures were investigated in the laboratory at 25 ± 1 °C and 65 ± 5% RH. More than 50% of T. pyri protonymphs preying on larvae of K. aberrans or E. finlandicus reached the adult stage. The mean developmental time of T. pyri from the protonymphal stage to adulthood was 6.3 days when feeding on K. aberrans and 7.5 days when feeding on E. finlandicus. The majority (approximately 90%) of K. aberrans protonymphs feeding on larvae of T. pyri or E. finlandicus died before reaching the deutonymphal stage; in both cases only one individual completed juvenile development. Euseius finlandicus larvae require food to reach the subsequent life stage, in contrast to larvae of T. pyri and K. aberrans, which usually do not feed at all: 10% of E. finlandicus immatures feeding on larvae of K. aberrans or T. pyri completed juvenile development (mean developmental time from larva to adult 7.0 and 6.7 days, respectively). Cannibalizing immatures of T. pyri and K. aberrans were able to reach adulthood, whereas those of E. finlandicus were not. Unfed immatures of T. pyri lived longer than the corresponding stages of E. finlandicus and K. aberrans. The present study indicates that phytoseiid immatures are suitable prey for developing stages of polyphagous phytoseiids. Since E. finlandicus, T. pyri and K. aberrans partly inhabit the same plant species, their immatures can be regarded as potential prey for competitive phytoseiids in times of food scarcity.     

A Genetic Analysis of Mancozeb Resistance in Typhlodromus Pyri (Acari: Phytoseiidae)

A field population of Typhlodromus pyri (Acari: Phytoseiidae) tolerant to mancozeb was selected in the laboratory. After 10 mancozeb selections the LC₅₀ value for mancozeb was 73 times higher in the selected-10 strain compared to the standard susceptible strain. A genetic analysis using reciprocal crosses and backcrosses of female F₁ progeny found no maternal effect. Resistance in the selected-10 strain was codominant in expression, dominance value was about −0.1. Backcrosses between F₁ females and the susceptible strain indicate that the resistance to mancozeb could be principally conferred by a predominant gene, but additional factors would also be involved.     

Infochemical-Mediated Niche Use by the Predatory Mites Typhlodromalus manihoti and T. aripo (Acari: Phytoseiidae)

In Africa, Typhlodromalus manihoti and T. aripo, two introduced predators of the cassava green mite Mononychellus tanajoa, occupy different parts of cassava foliage. In the present study, niche use by these two predators, as mediated by prey-induced infochemicals, was investigated. In response to prey feeding damage, cassava plant parts emit volatile blends, that attract phytoseiidae predators. When given a choice between old cassava leaves infested with M. tanajoa and either apices or young cassava leaves infested with M. tanajoa, T. aripo displayed a marked preference for odors emitted from either infested apices or infested young leaves over infested old leaves but showed no preference for odors from apices versus young leaves, all infested with M. tanajoa. Typhlodromalus manihoti did not discriminate between volatiles from the three infested cassava plant parts. Our data show that T. aripo uses differences in volatile blends released by infested cassava plant parts and restricts its fundamental niche to a realized niche, which enables coexistence with its competitor T. manihoti.     

Inter- and intraspecific predation on immatures by adult females in Euseius finlandicus, Typhlodromus pyri and Kampimo dromus aberrans (Acari: Phytoseiidae)

In a series of experiments, the interspecific predation and cannibalism on immatures by the adult females of Euseius finlandicus, Kampimodromus aberrans and Typhlodromus pyri were examined under laboratory conditions. The three species showed differing tendencies to prey on each other's motile immature stages. Euseius finlandicus females consumed more larvae and protonymphs than the females of T. pyri and K. aberrans. In cages without free water E. finlandicus ate a range of 6.51 larvae or 5.31 protonymphs of T. pyri and 5.27 larvae or 5.95 protonymphs of K. aberrans per female per day. Kampimodromus aberrans and T. pyri females exhibited a greater tendency to prey on heterospecifics than on conspecifics. When feeding on phytoseiid immatures and without free water, T. pyri females survived longer and laid more eggs than the females of E. finlandicus and K. aberrans. Adult females of E. finlandicus and T. pyri having free water and preying on heterospecific protonymphs were able to maintain egg laying during the whole experimental period of 12 days. The females of all three species had difficulties in piercing phytoseiid eggs, and the number of sucked eggs per female per day was low. Kampimodromus aberrans females ate 0.48 eggs of T. pyri daily, which was the highest recorded number. The great tendency to interspecific predation on motile immature phytoseiids by the females of E. finlandicus is discussed with regard to the dominance of this species on deciduous trees and bushes in Austria.     

Impact of vineyard pesticides on a beneficial arthropod, Typhlodromus pyri (Acari: Phytoseiidae), in laboratory bioassays

Laboratory bioassays were conducted to evaluate the effects of six vineyard pesticides on Typhlodromus pyri Scheuten (Acari: Phytoseiidae), a key predator of the mite Calepitrimerus vitis Nalepa (Acari: Eriophyoidae), in Pacific coastal vineyards. Materials tested were whey powder, 25% boscalid + 13% pyraclostrobin (Pristine), 40% myclobutanil (Rally), micronized sulfur (92% WP), 75% ethylene bisdithiocarbamate (mancozeb; Manzate), and 91.2% paraffinic oil (JMS Stylet), all applied at three concentrations. Pesticide dilutions were directly sprayed onto T. pyri adult females and juveniles, and each treatment was assessed to determine effects on direct mortality and fecundity. Five of the six pesticides tested resulted in < 50% mortality to adult and juvenile T. pyri for all concentrations 7 d after treatment. Paraffinic oil treatments displayed direct mortality > 50% for adult and juvenile assays and resulted in significantly higher mortality. Sublethal effects were more pronounced than acute pesticide toxicity, particularly in juvenile mite bioassays. Significant decreases in fecundity were detected in the sulfur and mancozeb treatments compared with the control in juvenile tests. The relative percentage of fecundity reduction for juvenile mites was highest when applying mancozeb (> 70%), sulfur (> 25%), or myclobutanil (> 20%). Adult mites displayed the greatest reductions in fecundity from applications of paraffinic oil (> 20%) or mancozeb (> 15%) treatments. Boscalid (+ pyraclostrobin) and whey displayed the least effect on fecundity across all bioassays. These results can be used to develop management guidelines in vineyard pest management practices to help conserve and enhance predatory mite populations.     

Jerking in predaceous mites (Acari: Phytoseiidae) with emphasis on larvae

Jerking, a behavior involving a pronounced and often repeated lunging of the body, was studied in larvae among seven species of phytoseiid mites, Euseius finlandicus, Galendromus occidentalis, Neoseiulus californicus, Neoseiulus fallacis, Phytoseiulus macropilis, Phytoseiulus persimilis and Typhlodromus pyri, and in the nymphal stages of N. fallacis. This behavior was observed in larvae of six of the seven species and in all active immature stages of N. fallacis. Jerking was usually triggered by direct contact with a con- or heterospecific mite, although jerking occurred occasionally without direct contact in N. fallacis larvae. The larval jerking tendency (the mean fraction of contacts resulting in jerking) of a species was significantly correlated with a tendency to congregate with conspecifics. Congregated larvae often probed their immediate area and each other with the front legs, repeatedly jerking in response to contact with neighboring larvae. Species with larvae having higher jerking tendencies also jerked more repeatedly per jerking event. Among species, there was no consistent relationship between jerking tendency and larval age. Larvae of N. fallacis had a higher tendency to jerk than protonymphs and deutonymphs, and they had both the highest jerking and congregating tendencies among larvae of all species studied. An individual was more likely to jerk when approached by another mite than when it initiated contact, and when observed in N. fallacis larvae that initiated contact, jerking was sometimes aggressive. Jerking N. fallacis larvae experienced fewer and shorter periods of probing by the mouthparts and fewer attacks from cannibalistic adult females than anesthetized larvae that were unable to jerk.     

Prey preference of predatory mite Amblyseius swirskii (Acari: Phytoseiidae) on Tetranychus urticae (Acari: Tetranychidae) and Bemisia tabaci (Hemiptera: Aleyrodidae)

Tetranychus urticae Koch (Acari: Tetranychidae) and Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae) are major pests in greenhouse crops. Recently, Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae) was shown to be an effective biological control agent of both pests. Therefore, the prey preference of A. swirskii was determined using immature stages of T. urticae and B. tabaci in three various treatments based on Manly's β preference index (β). These treatments consisted of immature stages of two prey species (egg, first and second instar nymphs) with densities 12:12, 6:6 and 3:3, respectively, and with 13 replicates. After 24?h starvation, same-aged females of A. swirskii were added to the leaf discs. All experiments were done on bean leaf discs in Petri dishes (8?cm in diameter) in laboratory conditions with 25?±?2°C, 70?±?5% relative humidity and the photoperiod of 16L:8D hours. Comparing the preference indices using t-tests indicates a significant preference of the predator on eggs (t?=?10.80, df?=?24, P?t?=?8.17, df?=?24, P?T. urticae than B. tabaci. Our findings suggest that developmental stages of prey have effect on the prey selection by A. swirskii.     

Side effects of mancozeb on Typhlodromus pyri (Acari: Phytoseiidae) in vineyards: results of multi-year field trials and a laboratory study

The side effects of mancozeb on the predatory mite Typhlodromus pyri were studied in 4-year field trials on grapevine and in the laboratory. In the field, the effect of mancozeb varied according to previous mancozeb use. In vineyards where mancozeb had commonly been used over years, this fungicide is generally slightly toxic, in some cases moderately toxic and rarely toxic. In plots were mancozeb has never been used, its effect on T. pyri was more pronounced and varied from moderately toxic to toxic. Despite the toxicity of mancozeb, T. pyri populations have never been eradicated. Laboratory results obtained with the French CEB guideline no. 167 confirmed those of the 4-year field study: mancozeb was significantly more toxic to T. pyri populations collected in plots where it had rarely been used before the field experiment. In plots where mancozeb had been used for a long time, the susceptibility of T. pyri populations to this fungicide was reduced and female survival, fecundity but also viability of female progeny were less affected by mancozeb. Even though toxicity of mancozeb increased in controlled conditions, a significant correlation was established between field and laboratory results.     

Temperature-related development and population parameters for Typhlodromus pyri (Acari: Phytoseiidae) found in Oregon vineyards

The beneficial mite Typhlodromus pyri is a key predator of grapevine rust mite Calepitrimerus vitis in Pacific coastal vineyards. Rust mite feeding has been associated with damage such as stunted, deformed shoot growth and reductions in fruit yield. The life history traits of T. pyri were assessed at seven constant temperatures (12.5, 15, 17.5, 20, 25, 30 and 35 °C) to determine population parameters providing data to better predict biological control of C. vitis populations by T. pyri in vineyards. Successful development from the egg to adult stage was observed at temperatures ranging from 15 to 30 °C. Constant exposure to 12.5 and 35 °C resulted in 100 % mortality in immature T. pyri. Developmental times, fecundity and longevity were highest at 25 °C. The estimated minimum and maximum developmental thresholds were 7.24 and 42.56 °C, respectively. Intrinsic rate of increase (r ( m )) was positive from 15 to 30 °C indicating population growth within this range of temperatures. Net reproductive rate and intrinsic rate of increase were greatest at 25 °C. These developmental parameters can be used to estimate population growth, determine seasonal phenology and aid in conservation management of T. pyri. Results presented in this study will aid in evaluating the effectiveness of T. pyri as a key biological control agent of C. vitis during different periods of the growing season in Pacific Northwest vineyards.     

Distribution of the Predatory Mite Typhlodromus pyri (Acari: Phytoseiidae) on Different Apple Cultivars

The effect of apple cultivar on the distribution of the predatory mite Typhlodromus pyri was studied in an experimental orchard where spider mites occurred at negligible densities. Seven apple scab-resistant cultivars, showing some differences in their leaf morphology, were considered. In particular, their leaf blade was classified according to four levels of pubescence. The distribution of T. pyri along the shoots was also studied. In the first experimental year the colonization of different cultivars by T. pyri showed definite patterns, sometimes influenced by the occurrence of eriophyids. One year later, T. pyri abundance again showed some differences among cultivars in conditions of prey scarcity. In both years large phytoseiid populations were recorded on the cultivar N.Y. 18491, despite the low eriophyid occurrence, probably because of its highly pubescent leaf undersurfaces. In contrast, Prima and TSR 29T219, characterized by slightly pubescent leaf undersurfaces, supported low phytoseiid densities independently of prey availability.     

Intra- and interspecific predation by adult female Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) when provisioned with varying densities and ratios of Tetranychus urticae (Acari: Tetranychidae) and phytoseiid larvae

The phytoseiid mites Metaseiulus occidentalis (Nesbitt) and Typhlodromus pyri Schueten are used together and alone as biological control agents against tetranychid pest mites of apple. Their effectiveness as control agents may be impacted by intraguild predation. The effects of prey species and prey density on the rates of inter- and intraspecific predation and oviposition by these two predators were investigated through a series of experiments. Adult female predators were given prey as mixed populations of phytoseiid larvae and larvae of a more preferred species, the spider mite, Tetranychus urticae Koch, at different densities and ratios. Typhlodromus pyri, more of a generalist predator, showed higher rates of predation and cannibalism on phytoseiid immatures at most prey densities and ratios. Manly preference indices indicated that T. pyri switched to feed on phytoseiid larvae at higher prey levels and ratios of T. urticae than M. occidentalis. This greater ability to use phytoseiid larvae as prey may help stabilize T. pyri populations when more preferred prey is unavailable. This may, in part, explain the observed persistence of T. pyri populations when M. occidentalis populations were decreasing in orchard test plots.