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1.
The development of effective biological control programs in which predators are integrated with other natural enemies such as parasitoids and entomopathogens requires an understanding of their interactions. In this study we examined the extent to which the omnivorous mirid bug Dicyphus hesperus, an effective biological control agent of greenhouse whiteflies, accepts prey that are either parasitized by the specialist whitefly parasitoid, Encarsia formosa or infected by the generalist fungal entomopathogen, Beauveria bassiana. In non-choice laboratory experiments, we measured how parasitism and infection of greenhouse whitefly, Trialeurodes vaporariorum, as related to parasitoid age and course of fungal infection, might alter the probability of feeding by second instar or adult female predators. Predation by D. hesperus was similar for parasitized and unparasitized whiteflies, regardless of parasitoid age. However, predators generally avoided feeding on infected whiteflies, particularly when infection was manifested through the presence of oosporein or hyphae on the surface of prey.  相似文献   

2.
Abstract This study examined the diel activity pattern and the effect of diel activity pattern on predation rate and prey finding of Dicyphus hesperus Knight (Heteroptera: Miridae). To determine the diel activity pattern of D. hesperus, starved females were placed on tomato leaflets Lycopersicon esculentum Mill. (Solanaceae) under zero, low, or high light intensities at 02:00, 08:00, and 14:00 h, respectively, and the amount of time spent walking or resting during a 30‐min interval was recorded. Predation rates of D. hesperus females on Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) egg patches were determined under either a L16:D8 (long day) or L8:D16 (short day) diel period. Egg patches were removed from D. hesperus females after either 8 or 16 h of dark or 16 or 8 h of light, and the number of eggs consumed was counted. Dicyphus hesperus females spent more time searching for prey at night than during the day. Females ate eggs at a higher rate during the night than during the day. Overall, D. hesperus females had higher predation rates when reared under a long day diel cycle compared with females reared under a short day diel cycle. More females reared at the L16:D8 diel cycle found the egg patch during the night than during the day. There was no difference in egg patch finding between night and day for females reared at L8:D16. Overall, L16:D8 reared females found more egg patches than females reared at L8:D16. Therefore, D. hesperus females are more active and find and consume prey at a higher rate at night than day.  相似文献   

3.
The effects of water stress (produced by water deprivation and prey feeding) on plant feeding were investigated in the omnivorous predator Dicyphus hesperus Knight (Hemiptera: Miridae). The objective was to determine if prey feeding aggravated water deficits and thus increased plant feeding. We measured plant feeding in a factorial experiment where female D. hesperus were prepared for experiments by providing or withholding water and/or prey for 24 h. We then evaluated the amount of plant feeding on Nicotiana tabacum seedlings by the direct observation of insects at three different densities of the prey, Ephestia kuehniella eggs. The amount of plant feeding, as measured by frequency of plant feeding bouts and time spent plant feeding during observation, was significantly greater for water‐deprived individuals than for those that had been provided with water. Individuals that had been provided with prey fed on plants at a significantly higher frequency than prey‐deprived individuals at two of the prey densities used in the experiment. These results support the hypothesis that plant feeding in zoophytophagous Hemiptera facilitates prey feeding by providing water that is essential for predation.  相似文献   

4.
1. Dicyphus hesperus Knight (Heteroptera: Miridae) nymphs were fed from egg hatch to the adult stage on Ephestia kuehniella eggs provided either alone or in combination with tomato leaves or with a supplementary water source. 2. Only 6% of individuals completed nymphal development on a diet of eggs alone. In contrast, a high proportion of nymphs completed development on a diet of eggs when either tomato leaves (97%) or a supplementary water source (88%) were provided. 3. The development times of nymphs given access to leaves were significantly shorter than those of nymphs given access only to supplementary water. 4. Adult female D. hesperus that were given access to tomato leaves prior to feeding trials consumed significantly more eggs in a 4‐h period than females that were dehydrated before trials. Dehydrated females that were allowed access to water for 3 h before trials consumed an intermediate number of eggs. 5. Plant feeding or access to some other water source is required for prey feeding, growth, and development in D. hesperus, and acquisition of water is proposed as a primary function of plant feeding. In addition, D. hesperus derives nutrients from plant feeding that increase the rate of nymphal development, although nymphs cannot complete development when provided only with tomato leaves. 6. Three simple models are presented of feeding behaviour in predatory Heteroptera where the amount of plant feeding either decreases, increases, or is constant as a function of the amount of prey feeding. The models are discussed with reference to the results and the probable multifunctional nature of plant feeding in predatory Heteroptera.  相似文献   

5.
An ongoing debate in biological control consists of whether interference between biological agents can disrupt pest control. This study investigated the outcome of interactions between the entomopathogen Beauveria bassiana with the whitefly predator Dicyphus hesperus and the parasitoid Encarsia formosa, as well as their effect on the control of the greenhouse whitefly Trialeurodes vaporariorum on greenhouse tomato crops. Our objective was to determine whether the generalist B. bassiana would disrupt biological control by interfering with D. hesperus or E. formosa. In experimental greenhouses, whitefly, parasitoid and predator populations were established, and over 27 days, tomato plants were sprayed with three applications of the B. bassiana based product BotaniGard® (5.13×103 conidia/mm2) or water (control). Populations of greenhouse whitefly and biological control organisms were regularly monitored in control and B. bassiana-treated compartments. Overall, 10.6% of all whiteflies in treated compartments were infected, and 0.98% were both infected and parasitized. There were 31.7 and 22.3% fewer immature and adult whiteflies, respectively, on B. bassiana-treated plants relative to controls. Parasitism by E. formosa and predation by D. hesperus occurred at rates of 7.5 and 2.5%, respectively, in B. bassiana-treated compartments, and 5 and 6%, respectively in control compartments. Our study suggests that applications of B. bassiana for short-term biological control of greenhouse whiteflies are compatible with the concurrent use of E. formosa and D. hesperus on greenhouse tomato crops.  相似文献   

6.
Intraguild predation of Orius majusculus (Reuter) (Heteroptera: Anthocoridae) on Encarsia formosa (Gahan) (Hymenoptera: Aphelinidae), both natural enemies of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), was studied under laboratory conditions. The experiments quantified prey consumption by 5th instar nymphs and adults of O. majusculus offered unparasitised 3rd, early 4th or 4th instar B. tabaci nymphs or parasitised nymphs containing 2nd or 3rd larval instar or pupal parasitoids. In addition, prey preference of the two stages of O. majusculus for parasitised or unparasitised whitefly nymphs was studied using nine different prey combinations. Both predator stages readily preyed upon on both unparasitised and parasitised B. tabaci. In no-choice experiments, predation on 3rd instar E. formosa by adult predators was the highest, while predator nymphs preyed most on unparasitised 3rd instar B. tabaci and 2nd instar parasitoids. Predation of predator stages was lowest on 4th instar B. tabaci and E. formosa pupae. In all prey combinations, both stages of O. majusculus showed a significant preference for parasitised over unparasitised whitefly nymphs except for the combination of 5th instars of O. majusculus with early 4th instar whiteflies and E. formosa pupae. The results indicate that intraguild interactions between O. majusculus and E. formosa may have negative effects on biological control of B. tabaci.  相似文献   

7.
Pesticides can negatively affect many life history traits of natural enemies. In this study, we studied the effects of three insecticides with different modes of action on the bionomics of Encarsia formosa, a parasitoid of the greenhouse whitefly, Trialeurodes vaporariorum (Westwood). Buprofezin (800 mg (a.i.)/L) and pyriproxyfen (50 mg (a.i.)/L) were selected among IGRs, while fenpropathrin (250 mg (a.i.)/L) was selected from the Pyrotheroids. Adults of E. formosa were treated via exposure to residues of insecticides on leaf discs. Our results revealed that buprofezin and pyriproxyfen did not affect longevity and fecundity, while fenpropathrin significantly reduced the longevity and fecundity of treated wasps. Results of logistic regression revealed that control, buprofezin- and pryproxyphen-treated E. formosa was a function of host density and followed a type II functional response. In contrast, E. formosa treated by fenpropathrin showed a type III functional response. Estimated attack rate for buprofezin and pyriproxyfen did not differ significantly from the control, whereas fenpropathrin-treated wasps showed a lower attack rate than the control. According to the obtained results, handling time of fenpropathrin-treated wasps was significantly higher (4.57 ± 0.5) than the control (2.83 ± 0.35). Our results showed that the maximum parasitism rate achieved by control wasps was 8.39, while the rate for buprofezin-, pyriproxyfen- and fenpropathrin-treated wasps was 6.99, 7.69 and 5.25, respectively. Overall, results suggest that buprofezin and pyriproxyfen can be used in an integrated pest management programme or biological control programme without destructive effects on the efficiency of this natural enemy in green houses.  相似文献   

8.
Searching behaviour of Encarsia formosa as mediated by colour and honeydew   总被引:2,自引:0,他引:2  
The habitat- and host-searching behaviour of female Encarsia formosa Gahan (Hymenoptera: Aphelinidae) was assessed using an airflow olfactometer and a filter paper test. Responses to different odour cues, colours, host-produced honeydew, non-host honeydew and single carbohydrates were determined. The parasitoid was not attracted to or arrested by odours emanating from clean tobacco leaves, tobacco leaves heavily infested with the host Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) and covered with honeydew, or honeydew alone. However, E. formosa females showed a significant response to green light transmitted through a tobacco leaf. The yellow part of the spectrum was partly responsible for this response. Thus, the long-range orientation is random with respect to the presence of hosts. Filter paper tests showed that the short-range searching behaviour is influenced by water soluble, non-volatile contact-kairomones contained in the host-produced honeydew. Contact with honeydew excreted by L3/L4 T. vaporariorum resulted in longer searching times than honeydew from adult T. vaporariorum or L3/L4 Bemisia tabaci Gennadius (Homoptera: Aleyrodidae). No difference was found between the response to honeydew excreted by adult and L3/L4 B. tabaci. The parasitoids' response to honeydew was unaffected by the host plant on which the whiteflies had fed. Non-host honeydew and single carbohydrates also affected the searching behaviour of E. formosa but to a lower extent than host honeydew. The possible differences in the carbohydrate and amino acid composition of the honeydew excreted by different life-stages of T. vaporariorum and B. tabaci are discussed.  相似文献   

9.
周淑香  李玉  张帆 《生态学报》2009,29(9):4732-4737
研究了持续高温(31℃恒温)和高适温交替(白天31℃/夜间25℃变温)条件对感染沃尔巴克氏体(Wolbachia)的丽蚜小蜂生殖和发育的影响.结果发现在持续高温条件下,丽蚜小蜂的生殖力和羽化率降低、寿命缩短、后代雄性比例增加,丽蚜小蜂从F1代开始产生雄性后代,且雄性比例在45%以上,F2代雄性比例达到最高值,表明高温冲击会对丽蚜小蜂的控害潜能产生不利的影响,因此在丽蚜小蜂的商业生产和田间应用中要注意环境温度,避免因高温带来的损失.在31℃恒温条件下连续培养3代或6代再转入25℃培养,丽蚜小蜂雄性后代比例和沃尔巴克氏体在丽蚜小蜂雌性群体中的感染率都能够很快恢复到高温处理前的水平.说明沃尔巴克氏体对环境变化有一个自动调节和适应的过程.变温条件下丽蚜小蜂从F2代开始产生雄性后代,但雄性后代比例较低(13%),表明短期高温冲击对丽蚜小蜂的生殖方式影响不大,但是短期高温冲击如果持续作用就会产生积累效应,影响丽蚜小蜂的后代性别.  相似文献   

10.
We investigated the spectral sensitivity and response to light intensity of Encarsia formosa (Hymenoptera: Aphelinidae), which is a key natural enemy of the greenhouse whitefly, Trialeurodes vaporariorum, and the tobacco whitefly, Bemisia tabaci (Hemiptera: Aleyrodidae). To do so, we used 15 monochromatic lights (emitting various specific wavelengths from 340 to 649?nm) and white light. E. formosa adults, which are diurnal insects, showed a positive phototaxis to a broad spectrum of light, with peaks of sensitivity at 414, 340, 450, and 504?nm. These results show that this parasitoid is generally more sensitive to short wavelength lights than long wavelength lights across all spectral ranges tested. Furthermore, E. formosa adults showed an increased phototactic response at low intensities and a decreased response at high intensities, for both ultraviolet light and violet light. Thus, E. formosa showed both colour and intensity preferences. This experiment provides a scientific basis for the development of colour traps for insect pest management and improves understanding of the ecological significance of colour vision by E. formosa.  相似文献   

11.
Interactions between natural enemies can be crucial for determining their overall control of pest species, yet the mechanisms that govern such interactions are often poorly understood. The risk of negative effects such as intraguild predation and the possibility of mitigating such risks are important components for ultimately determining the compatibility of biological control agents. We performed a group of experiments to determine whether the coccinellid Harmonia axyridis Pallas (Coleoptera: Coccinellidae) poses an intraguild threat to the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) and to see whether A. ervi is able to avoid predation by responding to the chemical tracks deposited by H. axyridis. We show that although H. axyridis does not readily consume A. ervi mummies, it preferentially consumes parasitized aphids over unparasitized aphids. We also show that A. ervi can defend against this threat by avoiding oviposition in the presence of H. axyridis chemical tracks. Aphidius ervi parasitized far fewer pea aphids Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae) on plants [Vicia faba L. (Fabaceae)] with H. axyridis chemical tracks in a no‐choice environment at a single‐plant scale. Similarly, when parasitoids could move freely between plants with and without tracks, A. ervi parasitism was higher on plants without tracks. Behavioral observations of A. ervi foraging suggested that this might be because of reduced A. ervi attack rates and patch residence times in the presence of H. axyridis tracks. Despite a risk of intraguild predation by H. axyridis, our study suggests that A. ervi may be able to mitigate this risk by altering its behavior in response to chemical cues.  相似文献   

12.
The native parasitoid Eretmocerus mundus Mercet and the predator Macrolophus pygmaeus Rambur are widely used to control Bemisia tabaci (Gennadius) in Mediterranean tomato greenhouses. An optimal biological control strategy for B. tabaci should take into account intraguild interactions between these natural enemies. In this study, predator's prey preferences and prey consumption were studied when offered different parasitoid and whitefly stages. The effect of the host plant on the adults of both natural enemies was also examined. M. pygmaeus preferred to consume B. tabaci over E. mundus when immature stages and adults of B. tabaci and E. mundus were offered. They consumed a larger amount of healthy B. tabaci nymphs and adults than of parasitised nymphs or E. mundus adults. The predator M. pygmaeus interfered with the reproduction of E. mundus females on cotton but not on tomato. However, B. tabaci nymphal mortality on tomato associated with parasitoid host feeding was also lower when the adult parasitoids coexisted with the predators. The joint release of M. pygmaeus and E. mundus adults did not increase the control of the whitefly B. tabaci.  相似文献   

13.
The compatibility of the encyrtid parasitoid Leptomastix dactylopii with the coccinellid beetle Cryptolaemus montrouzieri against the citrus mealybug, Planococcus citri, is determined by the extent of intraguild predation and interference by the predator. We tested the preference of the adults and fourth-instar larvae of C. montrouzieri for healthy mealybugs and parasitized mealybugs harboring 1-, 4-, 7- and 14-day-old parasitoid larvae. The experiments were conducted in no-choice (only unparasitized mealybugs or parasitized mealybugs of one parasitoid age were offered at one time) and choice (unparasitized and parasitized mealybugs of a specific age were offered simultaneously) tests. Both the adults and larvae of C. montrouzieri fed on unparasitized and parasitized mealybugs but strongly discriminated against the hardened mummies (14 days old). We also investigated the influence of the presence of C. montrouzieri to the foraging effectiveness of L. dactylopii. The level of parasitism by L. dactylopii was reduced from about 13 to 6% when the number of the C. montrouzieri was increased from zero to four. We recommended that the releases of C. montrouzieri should be conducted 14 days after the releases of L. dactylopii to reduce intraguild predation on the parasitized mealybugs and to avoid interference with the foraging parasitoids.  相似文献   

14.
The flight behaviour of Encarsia formosa Gahan (Hymenoptera: Aphelinidae), a parasitoid of Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) has been studied in response to the following natural components: bean plant (Phaseolus vulgaris L.) infested with T. vaporariorum (PHC, plant host complex), host damaged bean plant from which whiteflies were removed (HDP, host damaged plant), T. vaporariorum third instar larvae (H, host), uninfested bean plant (P, plant) and T. vaporariorum empty pupal cases (EPC, empty pupal cases). A combination of chemical and visual stimuli is necessary in host location from a distance by E. formosa and seems to deeply affect the dispersal of this parasitoid. The presence of host (PHC, H) or of its parts (EPC) led to a substantial increase of oriented flights suggesting the occurrence of host derived chemicals. At a closer range, a combination of visual and chemical stimuli deriving from all the components of the plant host complex elicited the transition flying-landing and the following landing on the source.  相似文献   

15.
Intraguild predation of the mealybug parasitoids Anagyrus pseudococci (Girault), and Leptomastix dactylopii Howard (Hymenoptera: Encrytidae) by Nephus kreissli Fürsch & Uygun (Coleoptera: Coccinellidae) was studied. The latter is a native predator of the important pest Planococcus ficus (Signoret) (Hemiptera: Pseudococcidae) on grapevines in Turkey. For this purpose, P. ficus of different ages parasitised by A. pseudococci or L. dactylopii, or by both A. pseudococci and L. dactylopii, were served to fourth instars and adults of N. kreissli as food. Experiments were conducted using two different treatments: no-choice (served unparasitised or only one stage of parasitised mealybug) and choice (served unparasitised and only one age of parasitised mealybug together), under controlled environmental conditions. Both fourth instars and adult predators were fed on two- and four-day-old mealybugs parasitised by A. pseudococci or on two-, four- and six-day-old mealybugs parasitised by L. dactylopii or by either A. pseudococci or L. dactylopii. The predators could not consume six-day-old mealybugs parasitised by A. pseudococci, eight-day-old mealybugs parasitised by L. dactylopii, or those parasitised by either of these parasitoids which had become mummified. While it was found that the adult predators preferred parasitised mealybugs to unparasitised, the larvae did not show a pronounced preference between parasitised and unparasitised mealybugs (except for mealybugs parasitised by A. pseudococci).  相似文献   

16.
昆虫群落中的集团内捕食作用   总被引:1,自引:0,他引:1  
与捕食、竞争等种间关系相比 ,集团内捕食作用是一种更为复杂的种间关系 ,广泛存在于各种生态系中。该文从它的概念、类型、作用特点、与其它种间关系的区别、存在的广泛性、发生强度的影响因素、化学信息调控及其对生物防治效果的影响等几个方面介绍了天敌群落中的这一典型的种间相互关系。根据参与者在集团内捕食作用关系中的利益地位的不同 ,可将集团内捕食作用分为单向性和双向性的 ,也可分成对称性的和不对称性的。参与者的生物学特性和各种环境条件对集团内捕食作用的发生程度和结果起着重要的调控作用。在多种天敌存在的系统中 ,集团内捕食作用影响着目标生物种群动态及其生物防治的效果。深入地研究这一种间关系对增强生物控害的功能有十分重要的意义  相似文献   

17.
We evaluated the influence of intraguild predation among generalist insect predators on the suppression of an herbivore, the aphid Aphis gossypii, to test the appropriateness of the simple three trophic level model proposed by Hairston, Smith, and Slobodkin (1960). We manipulated components of the predator community, including three hemipteran predators and larvae of the predatory green lacewing Chrysoperla carnea, in field enclosure/exclosure experiments to address four questions: (1) Do generalist hemipteran predators feed on C. carnea? (2) Does intraguild predation (IGP) represent a substantial source of mortality for C. carnea? (3) Do predator species act in an independent, additive manner, or do significant interactions occur? (4) Can the experimental addition of some predators result in increased densities of aphids through a trophic cascade effect? Direct observations of predation in the field demonstrated that several generalist predators consume C. carnea and other carnivorous arthropods. Severely reduced survivorship of lacewing larvae in the presence of other predators showed that IGP was a major source of mortality. Decreased survival of lacewing larvae was primarily a result of predation rather than competition. IGP created significant interactions between the influences of lacewings and either Zelus renardii or Nabis predators on aphid population suppression. Despite the fact that the trophic web was too complex to delineate distinct trophic levels within the predatory arthropod community, some trophic links were sufficiently strong to produce cascades from higher-order carnivores to the level of herbivore population dynamics: experimental addition of either Z. renardii or Nabis predators generated sufficient lacewing larval mortality in one experiment to release aphid populations from regulation by lacewing predators. We conclude that intraguild predation in this system is wide-spread and has potentially important influences on the population dynamics of a key herbivore.  相似文献   

18.
One of the factors that may complicate biological control of the greenhouse whitefly on Gerbera jamesonii by Encarsia formosa is the rosette shape of this ornamental, which differs from the vertical shape of most vegetable plants (cucumber, egg plant, tomato, etc.). Therefore, host-habitat location and the behaviour prior to landing on uninfested and infested leaves was studied. Attraction of E. formosa from a short distance by infested leaves could not be detected: the parasitoid females landed at random on uninfested and infested leaves. After the first landing, a redistribution of the wasps occurred on the leaves. After 24 h three times as many wasps were found on the infested leaves than on uninfested ones. In a dispersal experiment with four plants, E. formosa appeared to have no preference for landing on leaves of the medium age class, which is the age class on which most of the whiteflies in a suitable stage for parasitism occur. Twenty percent of the parasitoids were found on the plants 20 min after releasing them. These results were independent of the plant cultivar and the host density on the plants. In the course of 8 h, the number of E. formosa females recovered from plants increased linearly, and this increase was greater on plants where hosts were present and also greater on the plant cultivar with the lowest trichome density. After 24 h, the percentage of females was highest (56%) on plants with the highest host density. E. formosa females were arrested on leaves where hosts were present. Contrary to our expectation, the results from the two G. jamesonii cultivars that differed strongly in leaf hairiness were not significantly different in most experiments. Only at the high host density was parasitism found to be lower on the cultivar with the higher hair density. Parasitoids may walk on top of the `hair coverlet' of cultivars with high trichome density and, therefore, be hampered less than expected.  相似文献   

19.
In this research, for the first time, compatibility of imidacloprid and pyriproxyfen was assayed with both pupa and adult stages of Encarsia inaron Walker parasitizing Trialeurodes vaporariorum Westwood. First, the insecticides were sprayed on bean plants containing the parasitoid pupae, thenceforth, survival of the adult parasitoid was evaluated 24 h post-exposure to fresh, 1-, 4-, 7-, 14-, and 21-day-old residues of them on bean leaflets in Petri dishes. Both insecticides significantly reduce E. inaron adult emergence. Mortality of the parasitoid pupa in treatment with imidacloprid (69.7%) was significantly higher than mortality with pyriproxyfen (28.6%). Pyriproxyfen was non-toxic to the adult parasitoid when residues were dried in fresh and aged through the experiment, while in the same condition, imidacloprid significantly killed the adult parasitoids up to 7 days after application. Results have been discussed on the potential compatibility of the insecticides with E. inaron in an integrated pest management (IPM) approach.  相似文献   

20.
Dicyphine mirids are one of the most important groups of predators on tomato. In the Mediterranean region, several species in the genera Dicyphus, Macrolophus, and Nesidiocoris (Hemiptera: Miridae, Bryocorinae, Dicyphini) colonize protected horticultural crops. In Portugal, Nesidiocoris tenuis (Reuter) is increasingly abundant in the mirid species complex of tomato crops and appears to be displacing the native Dicyphus cerastii Wagner. In order to know whether intraguild predation (IGP) can explain the decreasing abundance of D. cerastii, we evaluated predatory interactions between adult females and first instars of D. cerastii vs. N. tenuis but also D. cerastii vs. Macrolophus pygmaeus (Rambur), as this species is also naturally present in horticultural crops in Portugal. Cannibalistic interactions were also tested for the same three species. All experiments were performed under laboratory conditions, in Petri dish arenas, in the presence or absence of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) eggs as alternative prey. Predation on both heterospecific and conspecific nymphs occurred only in the absence of alternative food. Intraguild predation was mutual and symmetrical between D. cerastii and M. pygmaeus. However, IGP was asymmetrical between D. cerastii and N. tenuis, favouring the first. Cannibalism was not significantly different among these mirid species. Our results show that D. cerastii has a greater capacity to feed on intraguild prey than N. tenuis. Therefore, IGP on small nymphs does not explain the abundance shift between D. cerastii and N. tenuis.  相似文献   

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