Pheromone-mediated optomotor anemotaxis and altitude control exhibited by male oriental fruit moths in the field

Abstract. In the field over short grass, pheromone-stimulated oriental fruit moth males, Grapholita molesta (Busck), flying under high windspeeds tended to steer courses more into the wind and to increase their airspeeds compared with those flying in low windspeeds.Thus, optomotor anemotaxis enabled the males to steer relatively consistent upwind track angles and to maintain an upwind progress of between c. 50–100 cm/s despite variable wind velocities.Zigzagging flight tracks were observed at both 10 m and 3 m from the source, as were tracks with no apparent zigzags.Transitions from casting to upwind flight or vice-versa were observed.The durations of the intervals between reversals during both upwind zigzagging flight and casting were consistent with those observed in previous wind-tunnel experiments.The control of altitude was more precise during upwind zigzagging flight than during casting.In general, the side-to-side deviations in the tracks were greater than the up-and-down deviations, with both the side-to-side and vertical distances and their ratios being consistent with previous wind-tunnel studies of pheromone-mediated flight.One difference between the field and laboratory flight tracks was that males in the field exhibited much higher airspeeds than in the wind tunnel.Males occasionally were observed to progress downwind faster than the wind itself, and further analysis showed that they were steering a downwind course in pheromone-free air following exposure to pheromone, which is the first time this has been recorded in moths.We propose that such downwind flight may aid in the relocation of a pheromone plume that has been lost due to a wind-shift, by enabling the moth to catch up to the pheromone as it recedes straight downwind away from the source.     

Appetitive flight patterns of male Agrotis segetum moths over landscape scales

An analysis is presented of the first harmonic radar studies of pheromone-plume locating flights of male Agrotis segetum moths over distances of up to 500 m. Upon release most moths flew in a direction having a downwind component. The first significant changes in flight orientations occur in the immediate vicinity of a pheromone source. Moths that were initially flying downwind change course and start flying crosswind whilst those that initially flew crosswind change course and start flying upwind. It is shown that such behaviour is consistent with the adoption of an effective plume-location strategy, and conditions are identified when downwind flights would be more advantageous than crosswind ones. Additionally, some of the complex flight patterns that can arise at later times are shown to be compatible with the adoption of an optimal biased scale-free (Lévy-flight) searching strategy. It is found that disruptive doses of sex pheromone can have a marked influence upon male moth flight patterns.     

Aim‐then‐shoot anemotaxis involved in the hopping approach of potato tuberworm moth Phthorimaea operculella toward a sex pheromone source

Male moths locate conspecific females by pheromone‐induced upwind flight maintained by detecting a visual flow, termed optomotor anemotaxis. Their behavioural pattern is characterized by an upwind surge in response to a pheromone stimulus and crosswind casting after odour loss, which is considered to be reset and restarted on receipt of another pheromone pulse. However, pheromone‐stimulated males of the potato tuberworm moth Phthorimaea operculella exhibit a series of short and straight intermittent flights, or hops, when moving upwind. It is unclear whether they navigate by employing the same behavioural pattern and wind detection mechanism as that used by flying moths. To analyze odour‐modulated anemotaxis in male potato tuberworm moths, a flat wind tunnel is constructed to give regular odour stimuli to an insect regardless of its location. Moths are subjected to pheromone pulses of different frequencies to test whether they show a behavioural pattern that is reset and restarted by a pheromone pulse. Moths on the ground are also subjected to crosswind shear to examine their detection of wind direction. Path analyses reveal that males surge upwind when they receive a pheromone pulse and exhibit casting by successive hops when they lose odour. This behavioural pattern appears to be similar to that of flying moths. When the direction of the airflow is switched orthogonally, males adjust their course angle accordingly when they are on the ground. It is suggested that, instead of optomotor anemotaxis, this ‘aim‐then‐shoot’ system aids the detection of wind direction, possibly by mechanosensory means.     

Migration to and dispersal from oilseed rape by the pollen beetle,Meligethes aeneus,in relation to wind direction

1 Host‐plant‐odour‐induced upwind anemotaxis is accepted as the main mechanism by which herbivorous insects find their host plant within an heterogenous environment, but field data supporting this hypothesis are scarce. 2 The flight directions of the pollen beetle Meligethes aeneus to and from a plot of winter oilseed rape and the direction of the wind were recorded concurrently. The beetles were sampled by eight double‐sided window traps encircling the plot. Distal sides of the traps, relative to the plot, sampled the beetles as they flew towards the plot, whereas the proximal sides of the traps sampled them as they flew away from the plot. Paired data on daily catch of beetles in distal or proximal traps and the volume of air impacting each trap were compared. 3 Correlations between daily catch of M. aeneus into distal traps and trap wind volume were negative, indicating that flights by both overwintered‐ and new‐generation insects towards the plot were upwind. 4 Correlations between daily catch of M. aeneus into proximal traps and trap wind volume varied with generation. Catch of overwintered‐generation M. aeneus was negatively correlated with trap wind volume, indicating that flight was upwind. Proximal trap catch of the new‐generation M. aeneus was weakly/modestly positively correlated with trap wind volume, indicating that flights were downwind or crosswind. 5 Understanding the effect of wind direction on flight of M. aeneus holds potential for improving the forecasting of their arrival and spatial distribution on the crop for integrated pest management.     

Behaviour of male pine sawflies, Neodiprion sertifer, released downwind from pheromone sources

This study investigated the behaviour of male European pine sawflies, Neodiprion sertifer Geoffroy (Hym., Diprionidae), that were released and observed downwind from pheromone traps baited with 100 g of the sex pheromone, (2S,3S,7S)-3,7-dimethyl-2-pentadecyl acetate. Releases were done at three distances; either at 5 m from one trap, or at 50 or 200 m from five traps, placed in a line perpendicular to the current wind direction. As control, males were released identically but without any pheromone source present. The behaviour of the males prior to take-off was studied. A total of 1729 males were released, and 80% of them took flight. Males took off significantly faster in the presence of pheromone. Grooming was significantly more frequent in presence of pheromone compared with control. In all pheromone experiments significantly more males displayed grooming, wing fanning and take-off towards the wind compared with the control. Weather data was simultaneously collected at the study site. Wing fanning was negatively correlated with wind speed. Grooming was not influenced by wind speed. Reduced levels of incoming short-wave radiation lowered the take-off frequency significantly. Pheromone-induced behaviour in diprionids seems to be less distinct than in other insects, e.g., Lepidoptera.     

Attraction range of a sex pheromone trap for the damson‐hop aphid Phorodon humuli (Hemiptera: Aphididae)

The attraction range of olfactory response by winged female gynoparae (autumn migrants that give birth to oviparae, the sexual females) and male damson–hop aphids Phorodon humuli (Schrank) is investigated in field experiments over 2 years by analyzing the spatial patterns of catches in concentric circles of yellow‐painted traps (60 in total) around a central trap releasing the species' sex pheromone, (1RS,7S,7aS)‐nepetalactol. Males are more likely than females to be found in the central trap, with 65.6% of the 1824 males caught there compared with 11.2% of 1346 females. Both morphs are more numerous in traps axial with the mean wind direction and centred on the pheromone‐release trap than at other angles. Males are approximately five‐fold more numerous in traps downwind than at similar distances upwind of the pheromone, showing that its presence stimulates landing. For males, the estimated active space of the lure extends 6 m downwind. Catches of females are equally numerous up and downwind of the pheromone lure because females orienting on the axis of the pheromone source continue to respond to visual cues in their flight path if they overshoot the olfactory one. For females, the active space of a pheromone lure is less than 2 m downwind. It is unimportant for either morph whether the pheromone‐release trap is yellow or transparent. In these experiments, both morphs orient with, track and probably arrive in the pheromone source trap from at least 26 m, the distance to the nearest aphid‐infested hops.     

Movement of the grape berry moth, Endopiza viteana: displacement distance and direction

Abstract. Mark–release–recapture is used to quantify displacement by adults of the North American grape berry moth, Endopiza viteana Clemens (Lepidoptera: Tortricidae) under field conditions. Moths marked with fluorescent dust are released eight times in the centre of a vineyard over 2 years, and recaptured using pheromone traps and interception traps. In vineyards, male moths are recaptured an average of 13.8 ± 0.8 m from the release site (maximum 58.2 m), whereas female displacement is similar with average flight distances of 11.4 ± 6.7 m (maximum 41.2 m). Increasing wind speed during moth flight activity periods suppresses displacement by both sexes, and females are less likely than males to fly in winds above 0.6 m s^?1. The majority of males are recaptured upwind from the release site or at a tangent to the overall mean wind direction when responding to pheromone traps, whereas female moths trapped in interception traps exhibit a large variability in direction from the release point. Releases of marked moths in woods adjacent to a vineyard demonstrates interhabitat movement by E. viteana males and by a single female. The average maximum displacement by males during interhabitat movement is 105.4 ± 3.9 m, significantly greater than the average maximum of 39.7 ± 6.7 m inside the vineyard habitat.     

Impact of (Z)-7-dodecenol and turbulence on pheromone-mediated flight manoeuvres of male Trichoplusia ni

Abstract. Turbulence and chemical noise are two factors which may influence pheromone-mediated flight manoeuvres of a moth in natural habitats. In this study, the effects of turbulence and the behavioural antagonist (Z)-7-dodecenol on flight manoeuvres of male Trichoplusia ni (Hübner) were evaluated in a wind tunnel. Male moths increase airspeed and course angles when turbulence is increased. This leads to significant increases in the length of flight tracks, but significant reductions in the time taken to reach a pheromone source. In less disturbed pheromone plumes, distributions of course angles and track angles of male T.ni show a prominent peak centred about 0° relative to the upwind direction, indicating that moths can temporarily steer directly upwind toward a pheromone source.
When (Z)-7-dodecenol is released 10 cm upwind of a pheromone source to form an overlapping plume downwind, course angles, airspeeds and ground-speeds of male T.ni are reduced significantly compared with those in uncon-taminated pheromone plumes. This results in a longer flight time to reach a pheromone source. The decrease in flight speed would decrease the rate of contact with filaments, and thereby perhaps allow the moth to detect uncon-taminated pheromone filaments independently from filaments containing the behavioural antagonist.     

Factors influencing adult male Grapholita molesta dispersal in commercial Malus and Prunus host crops

The oriental fruit moth (OFM), Grapholita molesta (Busck) (Lepidoptera: Tortricidae), has been an economic pest of apples (Malus spp.) and peaches (Prunus spp.; both Rosaceae) in the eastern USA since the 1930s. Successful management of G. molesta with insecticides requires careful timing of these products based on sex‐pheromone trap captures of male moths. In Pennsylvania, apple and peach orchards are often planted adjacently. Factors such as trap distance from release points, host crop, and wind were considered in relation to male moth movement within and between hosts. Our practical objective was to understand male G. molesta dispersal within and between these two hosts to achieve better management. Recapture of moths decreased over distance; the majority of moths were recaptured 50–100 m from the release point regardless of crop type. Male G. molesta released into commercial apple and peach orchards were most likely to remain in the crop of release and to not exhibit host shifts over the season, even after peach fruit were harvested. Our analyses of wind direction data with respect to recapture showed that the likelihood of recapture was similar regardless of the orientation of the traps relative to the mean wind direction at peak flight times. The apple and peach host crops appeared to affect male G. molesta dispersal to the extent that wind in orchards affects their perception of pheromone. Crop effects were most likely due to the traps closest to the release points being in the same crop as the release points.     

Dispersal ability in codling moth: mark–release–recapture experiments and kinship analysis

• 1 The present study focused on the dispersal patterns in the codling moth because such information is fundamental for determining the dynamics and genetics of the pest populations and for developing efficient management programmes.
• 2 We implemented mark–release–recapture experiments (MRR) with both male and female codling moths of two laboratory and one wild population using a sex pheromone and pear ester as attractants in delta traps. The experiments were conducted in apple orchards in central Greece over two consecutive years (2007–2008). In addition, kinship assignment tests were applied on 303 genotyped individuals (11 microsatellite loci) from two contiguous apple orchards in central Greece aiming to estimate the dispersal of fertilized females.
• 3 Both MRR and kinship analysis revealed that most male and female adults dispersed within 80 m, whereas some individuals moved at longer distances (maximum distance of approximately 200 m). A Bayesian analysis on microsatellite data revealed that exchange rates of codling moths between neighbouring orchards ranged among generations from 17.6 to 32.7%. The exchange rate between these orchards estimated by kinship analysis was 25.6% over all generations.
• 4 The collected data confirm the view of the sedentary nature of coding moth and indicate that genotypes able to migrate at long distances are not present in the studied area. The availability of food resources within orchards during the growing season is one possible factor that could favour this sedentary behaviour.

     

Distant olfactory response of the onion fly, Delia antiqua, to host-plant odour in the field

ABSTRACT. Distant olfactory orientation of female adult Delia antiqua (Meigen) to the host-plant Allium volatile dipropyl-disulphide (DPDS) was examined in the field using mark-release-recapture experiments and observations of flight behaviour. Onion-reared, post-diapause, virgin females from a laboratory colony dispersed upwind when released in the centre of 25, 50 and 100 m radius circles of eight 50 μl UDPDS baits. Percentage recapture and dispersal directedness did not decrease as a function of increasing distance to baits. In all cases the mean flight direction of recaptured flies closely correlated with mean wind direction. However, modes of the circular distributions of recaptured flies were located further crosswind when odour-baits were more distant. When distance was held constant (25 m) and DPDS concentration serially reduced (500–0.05 μ/bait), flies dispersed randomly in the absence of DPDS, crosswind in response to 0.05 μl baits and upwind in response to all other baits. Percentage recaptures on DPDS-baited traps of all concentrations were significantly greater than unbaited traps. Results from markrecapture studies were corroborated by observations of flight behaviour downwind. Flies located 100 m downwind from 50 μl DPDS baits flew upwind on take off while take-off flights in the absence of DPDS were random. Our data indicate that Allium volatiles like DPDS are involved not only in the acceptance phase of host-selection, but also in the first and probably most important stage when onion flies are initiating search long distances downwind. We conclude that D. antiqua orients to host-plants using olfactory cues from distances that should be classified as long-range ( sensu Kennedy , 1977     

Antennal mechanosensors mediate sex pheromone‐induced upwind orientation in the potato tuberworm moth

Males of the potato tuberworm moth Phthorimaea operculella (Lepidoptera: Gelechiidae) locate conspecific females by a series of short and straight flights, or ‘hops’. On the floor of a wind tunnel, P. operculella can change their heading angles in response to wind direction shift, suggesting that they detect wind direction and orient upwind when on the ground. It is unlikely that they navigate in wind by vision‐induced optomotor anemotaxis as in many flying moths. To investigate the mechanism of wind direction detection in this species, their orientation behaviour in response to pheromone pulses is observed in a wind tunnel. Intact male moths orient upwind even in complete darkness. After the flagellum of one antenna is amputated, male moths still successfully orient upwind. However, they fail to head upwind when the basal segments of both of their antennae are immobilized with glue. The ability to surge upwind is restored by removing the glue from the antennae. Thus, the results of the present study indicate that males of P. operculella orient upwind in response to mechanoreceptive cues from mechanosensory organs on their antennae. In Lepidoptera, two distinct anemotactic mechanisms of different sensory modalities appear to coexist: optomotor anemotaxis when airborne and the aim‐then‐shoot anemotactic system mediated by antennal mechanoreception when on the ground.     

Behaviour of male European corn borer, Ostrinia nubilalis Hübner (Lep.; Crambidae) towards pheromone-baited delta traps, bucket traps and wire mesh cone traps

Abstract:  Pheromone-baited traps are used to monitor the flight of European corn borer (ECB; Ostrinia nubilalis ). Traps of various designs are available: bucket traps, delta sticky traps and wire mesh cone traps. However, these traps are not all equally efficient and little is known about the reasons for this difference in efficiency. We investigated the behaviour of ECB males towards bucket traps and delta traps by means of observation in a wind tunnel. We also carried out observations and capture trials with delta traps and wire mesh cone traps in field conditions. Our laboratory studies showed that ECB males were not optimally attracted and were poorly captured by a pheromone baited bucket trap. Furthermore, they were shown to readily enter delta traps in the wind tunnel but were caught after more than three passages through this trap. Field studies showed that wire mesh cone traps captured approximately six times more ECB males than delta traps. Observation of the behaviour of ECB males showed that this difference was due to more efficient moth capture rather than greater attraction of the moth. In total, 31.5% of the 219 males observed close to wire mesh cone traps were caught, vs. 2.5% of the 520 males observed close to delta traps. This greater efficiency is due to a better capture rate of the attracted males by wire mesh cone traps than by delta traps.     

Factors influencing the landing of maleEpiphyas postvittana (Walker) exhibiting pheromone-mediated flight (Lepidoptera: Tortricidae)

The effects of changes in various visual and olfactory properties of a white card surface on the landing position of male Epiphyas postvittanaexhibiting pheromone-mediated flight were studied in a wind tunnel. Males landed predominantly at the most downwind position of a surface in line with the pheromone source, regardless of the strength of the source. The position on the surface that males landed was strongly influenced by visual factors. The landing position of males appeared to be influenced by visual cues along all three axes of the surface. Decreases in either the dimension horizontally perpendicular to the wind direction or the vertical dimension resulted in greater numbers of males landing farther upwind on the surface than the downwind edge. Visual changes in the axis along the wind direction also affected the position at which males landed. For example, when presented with two white card surfaces with a 4- cm gap between them, males tended to land on the downwind edge of the upwind surface (on which the source was located). When the gap was bridged with clear Mylar, the landing pattern was significantly different, with the greater proportion of males landing on the downwind surface. However, when Mylar was placed on the plexiglass floor of the tunnel (in addition to bridging the gap), the landing pattern on the surface was not significantly different from that on the two surfaces without the Mylar bridge. It is suggested that during the prelanding and landing phases of pheromone-mediated flight, male moths orient to visual features of the surface containing the pheromone source rather than to visual features of the source (conspecific female moth) itself.     

To what extent can the tiny parasitoid wasps,Eretmocerus mundus,fly upwind?

Body miniaturization in insects is predicted to result in decreased flight speed and therefore limited ability of these insects to fly upwind. Therefore, tiny insects are often regarded as relying on passive dispersal by winds. We tested this assumption in a wind tunnel by measuring the burst speed of Eretmocerus mundus (Mercet), a beneficial parasitoid wasp with body length <1 mm. Insects were filmed flying upwind towards a UV light source in a range of wind speed 0–0.5 m/s. The Insects flew towards the UV light in the absence and presence of wind but increased their flight speed in the presence of wind. They also changed flight direction to be directly upwind and maintained this body orientation even while drifted backwards relative to the ground by stronger winds. Field measurements showed that the average flight speed observed in the wind tunnel (0.3 m/s) is sufficient to allow flying between plants even when the wind speed above the vegetation was 3–5 folds higher. A simulation of the ability of the insects to control their flight trajectory towards a visual target (sticky traps) in winds show that the insects can manipulate their progress relative to the ground even when the wind speed exceeds their flight speed. The main factors determining the ability of the insects to reach the trap were trap diameter and the difference between insect flight speed and wind speed. The simulation also predicts the direction of arrival to the sticky target showing that many of the insects reach the target from the leeward side (i.e. by flight upwind). In light of these results, the notion that miniature insects passively disperse by winds is misleading because it disregards the ability of the insects to control their drift relative to the ground in winds that are faster than their flight speed.     

How Trichogramma parasitoids use moth sex pheromones as kairomones: orientation behaviour in a wind tunnel

Abstract The orientation behaviour of Trichogramma egg parasitoids (T.evanescens Westwood and T.pretiosum Riley) in response to the sex pheromone of their noctuid hosts {Mamestra brassicae L. and Heliothis tea (Boddie)) was investigated in a wind tunnel. Wasps were released on platforms which served as models of leaves, and were exposed to overhead light and an air stream that was either clean or loaded with host sex pheromone. The wasps exhibited: upwind anemotaxis, which was not affected by odour; odour-modulated positive phototaxis; and odour-induced inverse orthokinesis. Compared with clean air, residence times, walking times and path lengths on a platform were higher in pheromone-loaded air than in clean air. In pheromone-loaded air, walking velocity was reduced. During locomotion on a horizontal platform, net movement was upwind, regardless of the presence or absence of pheromone in the air. On a platform inclined at 45^o, anemotaxis appeared offset by positive phototaxis. If wasps were released on top of a glass rod above a platform, host sex pheromone caused wasps to land shortly after take-off. These results are used to explain higher rates of parasitism of moth eggs in pheromone-treated plots in earlier field experiments.     

Observations on the flight paths of the day-flying moth Virbia lamae during periods of mate location: do males have a strategy for contacting the pheromone plume?

1. To maximize the probability of rapid contact with a female's pheromone plume, the trajectories of male foraging flights might be expected to be directed with respect to wind flow and also to be energetically efficient. 2. Flights directed either upwind, downwind, or crosswind have been proposed as optimal strategies for rapid and/or energetically efficient plume contact. Other possible strategies are random and Lévy walks, which have trajectories and turn frequencies that are not dictated by the direction of wind flow. 3. The planar flight paths of males of the day-active moth Virbia lamae were recorded during the customary time of its sexual activity. 4. We found no directional preference in these foraging flights with respect to the direction of contemporaneous wind flow, but, because crosswind encompasses twice the possible orientations of either upwind or downwind, a random orientation is in effect a de facto crosswind strategy. 5. A crosswind preference should be favoured when the plume extends farther downwind than crosswind, and this strategy is realized by V. lamae males by a random orientation of their trajectories with respect to current wind direction.     

Observations on the orientation of tsetse flies (Glossina pallidipes) to wind-borne odours

ABSTRACT. Observations of the upwind flight of Glossina pallidipes Austen near a source of host odour show that in the absence of a visual target the insects tend to overshoot the odour source in fast, low flight. There is no sign of the crosswind 'casting' flight which characterizes the behaviour of moths under similar circumstances, except that a 180 turn is executed to bring the tsetse flies back to the vicinity of the odour source in downwind flight. This may be followed by a second overshoot and another 180 turn before the insects alight within a metre or so of the source. The results indicate that the orientation of tsetse flies to host odour may involve a step-wise approach to the odour source, providing an opportunity for assessment of wind direction when the insects are at rest between successive bursts of flight.     

Spring dispersal ofSitona lineatus: The use of aggregation pheromone traps for monitoring

The spring dispersal ofSitona lineatus L. (Coleoptera; Curculionidae) was investigated on a Danish farm.S. lineatus dispersed by flight in the early spring on sunny, calm days with temperatures above ca. 15°C. Two thirds of the population ofS. lineatus dispersed from perennial leguminous crops (clover and lucerne) in the first period of flight activity. The next dispersal did not occur until one month later despite several intermediate flight activity periods. The first period of dispersal occurred before the germination of the spring sown summer host crop,Vicia faba L. The field bean crop was infested in three later invasions during a period of more than three weeks. The aggregation pheromone, 4-methyl-3,5-heptanedione, had a significant effect on captures of both males and females in cone traps placed on the ground. There was no effect of the pheromone on captures in yellow sticky traps placed 1.5 m above ground. The pheromone effect is discussed in relation to behavioural observations. Both types of traps may be used in a survey system for monitoring spring dispersal ofS. lineatus and optimal timing of insecticide spraying. However, the pheromone cone traps were highly specific whereas all kinds of flying insects were caught in the yellow sticky traps, thus making the latter traps less suitable for monitoring.     

Pheromone‐trapping the nun moth,Lymantria monacha (Lepidoptera: Lymantriidae) in Inner Mongolia,China

The nun moth, Lymantria monacha L., is one of the most important defoliators of Eurasian coniferous forests. Outbreaks during 2011–2015 in the natural/planted larch, and larch‐birch mixed forests of the Greater Khingan Range in Inner Mongolia, China, caused tremendous timber losses from severe defoliation and tree mortality. A series of trapping experiments were conducted in these outbreak areas to evaluate the efficacy of a synthetic species‐specific pheromone lure based on the female pheromone blend of European nun moth populations. Our results clearly show that the nun moth in Inner Mongolia is highly and specifically attracted to this synthetic pheromone, with few gypsy moths (Lymantria dispar) captured. Flight activity monitoring of L. monacha male moths using pheromone‐baited Unitraps at 2 locations during the summer of 2015 indicated that the flight period started in mid‐July, peaking in early August at both locations. Based on male moth captures, there was a strong diurnal rhythm of flight activity throughout the entire scotophase, peaking between 22:00 and 24:00. Unitraps and wing traps had significantly and surprisingly higher catches than the gypsy moth traps. Unitraps fastened to tree trunks 2 m above ground caught significantly more male moths than those at the ground level or at 5 m height. Male L. monacha moths can be attracted to pheromone‐baited traps in open areas 150–200 m distant from the infested forest edge. Our data should allow improvement on the performance of pheromone‐baited traps for monitoring or mass‐trapping to combat outbreaks of this pest in northeastern China.