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  • Monthly, Founded in 1950
    Supervisor:Chinese Academy of Sciences
    Sponsor:Institute of Zoology,Chinese Academy of Sciences
    The Entomological Society of China
    Domestic postal code: 2-153
    Foreign issuance code: Q61
    ISSN 0454-6296
    CN 11-1832/Q
Table of Content
20 October 2012, Volume 55 Issue 10
For Selected: View Abstracts Toggle Thumbnails
    Separation and identification of haemolymph proteins involved in immune response to Nosema pernyi infection in Antheraea pernyi (Lepidoptera: Saturniidae) larvae
    JIANG Yi-Ren, SONG Jia, QIN Yu-Lin, WANG Yong, ZANG Min, ZHONG Liang, YANG Rui-Sheng, SHI Sheng-Lin, DUAN Yu-Xi, QIN Li
    2012, 55(10):  1119-1131. 
    Abstract ( 2903 )   PDF (13806KB) ( 1716 )     
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    This study was aimed to preliminarily identify the proteins involved in the early immune response in the haemolymph from Antheraea pernyi larvae infected by Nosema pernyi. The 5th instar female larvae of A. pernyi were fed with N. pernyi spores and the proteins in the haemolymph were separated by SDS-PAGE, and the proteome profiles of the differential protein bands were further analyzed and identified by LC-MS/MS and proteomics analysis. Onedimensional SDS-PAGE analysis revealed that protein bands at 44 kD (AP44) and 28 kD (AP28) were enriched in the haemolymph of the female larvae at 144 h after feeding with the spores of N. pernyi. A total of 117 non-redundant proteins were identified by LC-MS/MS and searching the COG database, including 52 unique proteins from AP28, 53 unique proteins from AP44, and 12 proteins from both of them. Among them, 29 proteins are immune-related, including 15 proteins from AP28, such as heat shock proteins, ubiquitin-like proteins, ubiquitin-conjugating enzyme E2, juvenile hormone epoxide hydrolase, microtubule-associated protein RP/EB family member 3, lysozyme-like protein, ADP-ribosylation factor, putative defense protein and peptidoglycan recognition protein-like protein, and 10 proteins from AP44, such as DRK, prophenoloxidase and hemolin. Four proteins were identified from both AP28 and AP44, including heat shock protein hsp21.4, prophenoloxidase and basic attacin. The results provide the basis for further study on the mechanism of immune response and defense against N. pernyi infection in A. pernyi.
    Cloning and developmental expression of dynactin p62 gene in the Chinese honeybee, Apis cerana cerana (Hymenoptera: Apidae)
    ZENG Jing, PAN Qi-Zhong, WANG Zi-Long, WU Xiao-Bo, YAN Wei-Yu, ZENG Zhi-Jiang
    2012, 55(10):  1132-1141. 
    Abstract ( 2821 )   PDF (15470KB) ( 1493 )     
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    To explore the expression characteristics of dynactin p62 gene in the Chinese honeybee, Apis cerana cerana, the genomic DNA sequence (GenBank accession no. JX101463) and mRNA sequence (GenBank accession no. JX101464) of dynactin p62 in A. cerana cerana were cloned, and a quantitative analysis of its expression level at different developmental stages of the queen, worker and drone (3 and 6 dold larvae, and newly eclosed bees) was conducted using realtime PCR. The full-length genomic DNA and mRNA sequences of A. c. cerana dynactin p62 are 2 403 bp and 1 491 bp, respectively, encoding 496 amino acids, and the predicted MW and pI are 56.49 kD and 8.31, respectively. Phylogenetic analysis indicated that dynactin p62 of both A. c. cerana and A. mellifera gathered in the same clade. The A. c. cerana dynactin p62 transcript was clearly detected at different developmental stages of the queens and workers, and it was expressed significantly higher in newly eclosed adults than in larvae (P<0.05); and its expression was higher in workers than in queens (P<0.05), but with no obvious regularity in drone. These results suggest that dynactin p62 may be involved in caste differentiation in A. c. cerana.
    Genetic diversity of Beauveria bassiana in different silkworm populations from Guangdong and Guangxi, southern China, based on SSRs
    LU Si-Xing, LIU Ji-Ping, TANG Li, LI Xiang-Lin, MI Hong-Xia, YANG Ji-Long, YAN Yu-Wei, XU Shu-Lan
    2012, 55(10):  1142-1148. 
    Abstract ( 3130 )   PDF (3396KB) ( 1167 )     
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    The genetic diversity of 13 Beauveria bassiana strains in different silkworm (Bombyx mori) populations from Guangdong and Guangxi, southern China, was analyzed based on simple sequence repeats (SSRs). A total of 137 fragments were obtained with 9 SSR primers, on average 15 fragments each primer. Further analysis with POPGEN indicated that the inner-population genetic differentiation (Gst) was 0.0590, the polymorphic rate(PPL) was 97.73%, the Nei’s genetic diversity index (H) was 0.1896, and the Shannon’s information diversity index (I) was 0.3165. The level of genetic variation of Guangdong populations was higher (PPL=68.18%, H=0.1910, I=0.3044), while that of the Guangxi populations (PPL=65.91%, H=0.1713, I=0.1791) is smaller. The result showed that the genetic diversity of B. bassiana Guangdong strains was higher than that of B. bassiana Guangxi strains. The dendrogram of B. bassiana Guangdong and Guangxi strains was constructed by UPGMA cluster analysis based on the Nei’s genetic distance, and it is found that different strains from different collecting localities are divided into two clades. The results indicate that the genetic structure of B. bassiana from Guangdong and Guangxi is very complicated, and there is certain degree of similarity between the strains of Beauveria from the partial silkworm region, suggesting that it is feasible to trace the origin of B. bassiana with SSRs.
    The endosymbiotic microbiota of two geographical populations of the Asian citrus psyllid, Diaphorina citri, in China and its ability of pathogen transmission
    MA Xiao-Fang, ZHANG Xue-Chao, XU Hai-Jun
    2012, 55(10):  1149-1153. 
    Abstract ( 2787 )   PDF (787KB) ( 1264 )     
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    Huanglongbing (HLB) is one of the most severe citrus diseases, which is usually transmitted by the Asian citrus psyllid (Diaphorina citri), and causes great economic loss. To understand the endosymbiotic microbiota of the citrus psyllid from Hainan and Fujian provinces in China, the partial conserved region of 16S rRNA was amplified by PCR, and subsequently the infection rate of each endosymbiont from two geographical populations of the citrus psyllid was detected by PCR using specific primer pairs in this stduy. Moreover, we detected the acquisition efficiency of the pathogen by adult citrus psyllid reared on Citrus reticulata cv. Tankan infected with Candidatus Liberibacter asiaticus (CLas), and the transmission efficiency to Citrus reticulata cv. Subcompressa via CLas-positive psyllids. The results demonstrated that the two geographical populations of the citrus psyllid had the same composition of endosymbiotic microbiota, including α-Proteobacteria, Wolbachia spp., γ-Proteobacteria, mycetocyte symbionts, β-Proteobacteria, Oxalobacter, and β-Proteobacteria, Herbaspirillum, and the infection rate of each endosymbiont from the two geographical populations was more than 95%. After the adult citrus psyllids were reared on CLas-infected C. reticulata cv. Tankan for 28 d, the acquisition efficiency was as high as 82%. The whole tree of C. reticulata cv. Subcompressa was infected after rearing the CLas-positive psyllids on it for 75 d. The results would provide some theoretical basis for the study of citrus psyllid and the effective control of HLB.
    Evaluation of the toxicity of Cry1Fa to the Cry1Ac-resistant cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)
    WEI Ji-Zhen, GUO Yu-Yuan, GAO Xi-Wu, ZHANG Tao, LIANG Ge-Mei
    2012, 55(10):  1154-1160. 
    Abstract ( 3711 )   PDF (2644KB) ( 1080 )     
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    Cry1Fa-expressing corn and cotton have been commercially planted in U.S in order to effectively control several Lepidoptera pests. Making clear the cross-resistance of cotton bollworms (Helicoverpa armigera) to Cry1Ac and Cry1Fa and the synergism between the two toxins could provide the theoretical bases for rationally applying Cry1Fa+Cry1Ac cotton. The toxicity of Cry1Fa to the susceptible strain (96S) and the resistant strain (BtR, selected by Cry1Ac, the resistance ratio was 2 194.15-fold) was tested in our study. The results showed that the toxicity of Cry1Fa was lower than Cry1Ac to 96S, the LC50 value of Cry1Fa was 504.80 times as high as that of Cry1Ac, and there was 19.98-fold cross-resistance to Cry1Fa in the Cry1Ac-resistant strain (BtR). Cry1Fa mixed with Cry1Ac could increase the toxicity of Cry1Fa against 96S, and especially when the concentration of Cry1Fa was low, the synergistic effect was significant. However, the toxicity of Cry1Ac could be improved just when being added with high-dose Cry1Fa. Because the BtR strain had resistance to Cry1Ac, the toxicity of Cry1Ac to BtR reduced obviously. Mixing Cry1Fa into high-dose Cry1Ac could increase the mortality of BtR (P=0.0015, F=6.88, df=6), but the highest mortality was only 58.33%. The results of the 2-D-optimum orthogonal design experiment showed that Cry1Ac had significant effect to control the susceptible cotton bollworm (t1=13.76﹥t0.05), and the influence of interactive effect between Cry1Ac and Cry1Fa on toxicity was also significant (t22=2.42﹥t0.05t11=6.95﹥t0.05t12=3.43﹥t0.05). Cry1Ac and Cry1Fa had significant effect against the resistant cotton bollworm (t1=3.03﹥t0.05t2=2.59﹥t0.05), but Cry1Ac was the key determinant for the mortality of the susceptible and resistant cotton bollworms. The optimum concentration ranges of Cry1Ac and Cry1Fa were 1.41-2.10 μg/cm2. However, the interactive effect between Cry1Ac and Cry1Fa was not significant in the BtR resistant strain. Thus, although the crops expressing Cry1F+Cry1A can expand the spectrum of killing insects, however, because of the cross-resistance and the undesirable effect in managing the Cry1Ac-resistant bollworm, transgenic Cry1F+Cry1A cotton is not recommended for planting in China.
    Sublethal effects of chlorantraniliprole on the experimental populations of the white-backed planthopper, Sogatella furcifera (Hemiptera: Delphacidae)
    YANG Hong, WANG Zhao, JIN Dao-Chao
    2012, 55(10):  1161-1167. 
    Abstract ( 3834 )   PDF (1086KB) ( 1216 )     
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    In order to evaluate the effect of chlorantraniliprole, an insecticide in control of Lepidoptera larvae on rice plant, on the white-backed planthopper, Sogatella furcifera (Horváth), the rice-stem dipping method was used to test the toxicity of chlorantraniliprole to the experimental populations of S. furcifera, and the sublethal effects of chlorantraniliprole on S. furcifera were studied with life table. The results showed that the LC50 doses of chlorantraniliprole to the 3th instar nymphs and adults of S. furcifera were 19.26 mg/L and 19.69 mg/L, respectively. After the 3th instar nymphs of S. furcifera were treated with the LC10 and LC25 of chlorantraniliprole, respectively, both the life span and the fecundity per female decreased in F0 and F1, and significant difference existed between the treatment of LC25 and the control in F0 and F1 (P<0.05). Compared to the control, the treatment of LC25 caused a decrease of 52.94 and 78.45 eggs laid per female in F0 and F1, respectively, and 1.80 d and 2.62 d less in the life span of female in F0 and F1, respectively. The developmental duration of different stages were affected after S. furcifera nymphs were treated with the LC25 and LC10 of chlorantraniliprole. According to the survival rate and fecundity of S. furcifera treated by different doses of chlorantraniliprole, the life table of fecundity was constructed. We found that for LC10 and LC25 treatments the intrinsic rate of increase (rm) decreased by 11.25% and 34.41%, respectively, and the net reproductive rate (R0) decreased by 36.56% and 74.57%, respectively, while the mean generation time (T) and population doubling time (t) were delayed. The results suggest that LC25 and LC10 of chlorantraniliprole can decrease the population growth of S. furcifera, and this insecticide will restrain the natural population growth of S. furcifera when it is used in the field in controlling the Lepidoptera larvae on rice plants.
    Effects of temperature on development, reproduction and population growth of the western flower thrips, Frankliniella occidentalis (Thysanoptera: Thripidae)
    ZHANG Zhi-Jun, ZHANG You-Jun, XU Bao-Yun, ZHU Guo-Ren, WU Qing-Jun
    2012, 55(10):  1168-1177. 
    Abstract ( 4811 )   PDF (3014KB) ( 1390 )     
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    The western flower thrips, Frankliniella occidentalis (Pergande), is an invasive pest in China. Temperature is a key determinant for the stable establishment of thrips population in a new environment. In order to know the effects of temperature on the population growth of the western flower thrips, we investigated the development, survivorship and reproduction of F. occidentalis at different temperatures (15, 20, 25, 30 and 35℃), and calculated the parameters of population increase under different temperature conditions. The results indicated that the survivorship of F. occidentalis from egg-hatch to adult was the highest (62.8%) at 20℃, but no individual developed into adult at 35℃. The development rate increased significantly with rising temperature, the average developmental duration from egg to adult stage was about 30 d at 15℃, while the shortest was about 10 d at 30℃. The adult longevity was evidently shortened with rising temperature, at 15℃ the average lifespan was 36 d, the longest adult longevity even reached more than 60 d, while at 30℃ the shortest was only 10 d. The differences of fecundity (total number of the 1st instar nymphs produced by individual female) were not significant among 15, 20 and 25℃, with the values of 37.70, 32.56 and 37.80, respectively, which were all higher than that of the thrips at 30℃ (9.36 1st instar nymphs produced by individual female). The parameters of population growth [the intrinsic rate of increase (rm) and the net reproductive rate (R0)] were the highest at 25℃, with the values of 0.178 and 20.10 d-1, respectively, while those at 15℃ were only 0.096 and 18.67 d-1, respectively. From these results we concluded that temperature has great effect on the development of F. occidentalis, and the most suitable temperature range for this pest is 20-25℃, during which the intrinsic increase rate is higher. The development threshold temperature of F. occidentalis was estimated to be 7.4℃ and the effective accumulated temperature for development was 208.0 degreedays. In South China, Central China, North China and Northeast China, the estimated numbers of generations per year are 24-26, 16-18, 13-14 and 1-4, respectively, without consideration of other factors influencing the development and growth of the thrips. Especially in Kunming and Lijiang, Yunnan in Southwest China, the estimated number of generations per year are 13-15 and 8-10, respectively.
    Effects of infestation of Liriomyza huidobrensis (Diptera: Agromyzidae) on the contents of main nutrients, secondary metabolites and chlorophyll in cucumber leaves
    SUN Xing-Hua, ZHOU Xiao-Rong, PANG Bao-Ping, MENG Qing-Jiu
    2012, 55(10):  1178-1184. 
    Abstract ( 3135 )   PDF (1063KB) ( 1198 )     
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    The pea leafminer, Liriomyza huidobrensis (Blanchard), is a polyphagous insect pest on vegetables and ornamental plants. In order to investigate the interaction between this pest and its host plants, we assayed the contents of soluble sugars, soluble proteins, tannins, flavones and chlorophyll in cucumber leaves infested by the 1st to 3rd larvae of L. huidobrensis with different damage levels using anthrone colorimetric method, Coomassie brilliant blue staining, phosphor-molybdenum acid-phosphor-tungstenic acid colorimetry, Soxhlet extraction and acetone spectrophotometry, respectively. The results showed that the contents of soluble sugars, soluble proteins and chlorophyll in cucumber leaves decreased while those of tannins and flavones increased with the increase of damage by the larvae. The largest decrease amplitudes of the former three were 62%, 35% and 40%, respectively, and the highest increase amplitudes of the latter two were 26% and 53%, respectively. The influences of the larval infestation on the above compounds were systemic. The results suggest that the larval damage caused the decrease of nutrient contents and phytosynthesis of host plants while the increase of secondary metabolite contents, and therefore induced the resistance to herbivores. This study may provide a useful reference to further research on the interactive mechanisms between the leafminer and its host plants.
    Effects of different LED light sources on the behavior of Brithys crini (Lepidoptera: Noctuidae)
    TU Xiao-Yun, CHEN Yuan-Sheng, Chen-Juan, HU Zhang-Long, JIN Yi-Qin, XU Fei
    2012, 55(10):  1185-1192. 
    Abstract ( 3023 )   PDF (7272KB) ( 1223 )     
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    To understand the effects of different LED light sources on the behavior of Brithys crini (Fabricius), thanatosis, walking, resting, feeding, defecating, watching and other behaviors of larvae, mating and oviposition of adults in 4 different LED light sources (red light: 620-625 nm; green light: 520-523 nm; blue light: 465-467 nm; and white light: 460-465 nm) were observed under the laboratory conditions. The results indicated that: (1) there was significant difference in walking, resting and feeding time budget of larvae in different light (P=0.000 for walking, resting and feeding). The walking budget time in red and white light accounted for 98.06% and 61.16% of the total action time, respectively, the feeding budget time in green light accounted for 83.65%, and the walking and resting budget time in blue light accounted for 91.56%. There was significant difference in frequencies of walking, resting, feeding and watching. The walking frequencies in red and white light are the highest (47.37% and 32.00%, respectively), the feeding frequency in green light is the highest (40.00%), and the walking frequency in blue light is the highest (32.35%). (2) There was no significant difference in the thanatosis frequency of fallen larvae in different light, but there was significant difference in the thanatosis duration, and the duration in red light is longer than that in blue and white light. (3) There was no significant difference in eggs laid per mated or unmated female in different light. (4) From the 1st to 2nd day, when red, green, blue and white light was used in the dark phase, the peaks of mating and oviposition occurred in the dark phase of the 3rd day. When no light was used in the dark phase, however, the peaks occurred in the dark phase of the 1st day. The results suggest that there are different effects of different LED light sources on the behaviors of B. crini.
    Effects of different evolution models on DNA barcoding evaluated with the Noctuidae from Wuling Mountain, Hebei, northern China
    CHI Mei-Yan, HAN Hui-Lin, GAO Qiang, YANG Cong-Hui, JIN Qian, LI Jun, CHEN Fu-Qiang, WU Chun-Sheng, ZHANG Ai-Bing
    2012, 55(10):  1193-1204. 
    Abstract ( 3354 )   PDF (11136KB) ( 1810 )     
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    The effects of evolution models on success rates of species identification via DNA barcoding were evaluated based on specimens of 44 Noctuidae species collected from Wuling Mountain, Hebei, northern China. Different phylogenetic tree reconstruction methods, including the neighbor-joining, the maximum parsimony, the maximum likelihood and the Bayesian inference, were used to reconstruct phylogenetic trees based on the COI gene sequences of these species, and success rates of different models (12 models for the neighborjoining, 7 models for the maximum likelihood method, and 2 models for the Bayesian inference) were evaluated systematically. The results showed that there were no significant differences in success rates of species identification among 12 different models for the neighborjoining method, but significant differences in success rates were found among different evolution models for both maximum likelihood and Bayesian methods. As a comparison, the maximum parsimony method which is not based on evolution models obtained stable success rates of species identification. The 6 evolutionary models shared by the neighborjoining and maximum likelihood methods obtained different success rates between these two methods. In addition, we found that singletons in the database largely reduced success rates of species identification, suggesting that deep intraspecific sampling be required in the future studies of DNA barcoding.
    Phylogenetic analysis of genera of the tribe Blaptini based on the characteristics of defensive glands (Coleoptera: Tenebrionidae)
    LIU Chao, Ren-Guo-Dong
    2012, 55(10):  1205-1220. 
    Abstract ( 2954 )   PDF (13853KB) ( 6541 )     
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    The phylogenetic systematics of 65 species of Blaptini (Coleoptera: Tenebrionidae) belonging to 16 genera (including 7 new recorded genera) was dealt with based on the structure of defensive glands. The characteristics of the glands at the genera and tribal level including gland position, shape, length, width, distance between the two glands, thickness of glands, surface fingerprint whorl and wrinkles were summarized. The phylogenetic relationship of the 16 genera was established using SPSS19.0 and Hennig86 program (version 1.5). The result showed that the evolutionary relationship should be as follows: Prosodes>Blaptogonia>Tagonoides>Thaumatoblaps>Caenoblaps>Agnaptoria>Asidoblaps>Coelocnemodes>Dila>Gnaptor>Blaps>Pseudognaptorina> Nalepa>Belousovia>Gnaptorina>Itagonia. The result supports the geographic distribution feature of this group. The 16 genera of Blaptini were clearly separated based on the phylogeny of defensive glands’ morphological data, suggesting that these characteristics have important value in taxonomy.
    Progress in the molecular mechanisms of the innate immune responses in insects
    ZHANG Ming-Ming, CHU Yuan, ZHAO Zhang-Wu, AN Chun-Ju
    2012, 55(10):  1221-1229. 
    Abstract ( 3684 )   PDF (5441KB) ( 4396 )     
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    Insects lack acquired immune system processed by vertebrates, and rely solely on developed innate immune system to defend against the infection from alien pathogens such as bacteria, fungi, or virus and so on. In this article, we review the progress in the molecular mechanisms of the development and action of the innate immune responses in insects, especially focusing on the roles and functioning mechanisms of some important immunerelated molecules. The innate immune response in insects is divided into humoral immunity and cellular immunity. They function together to kill and eliminate the pathogens via phagocytosis, nodulation, encapsulation, coagulation, and melanization. When the insects are infected by the alien pathogens, the pattern recognition proteins/receptor (PRPs) in insects would firstly recognize and combine with pathogen-associated molecular pattern (PAMPs) in the pathogens, and then initiate the activation and regulation of a series of innate immune response containing immune-related serine proteases and serine protease inhibitors, and finally trigger the production of the immune effectors, such as antimicrobial peptides, melanin and so on, to kill the alien pathogens. The antimicrobial peptides are a kind of cationic peptides with low molecular weight, and have a broad spectrum of antimicrobial activity. They are produced via various mechanisms based on the types of invading pathogens. There are two signal transduction pathways regulating the production of antimicrobial peptides: Toll pathway activated by fungi and most of gram-positive bacteria, and Imd pathway initiated by gram-negative bacteria. These two pathways are contributed to the expression of different antimicrobial peptides due to the activation of different transcription factors.
    Age-stage two-sex life tables of the experimental population of Panonychus ulmi (Acari: Tetranychidae) on apples Malus sieversii subsp. kirghisorum and M. domestica Golden Delicious
    YIN Wan-Dong, YAN Wen-Tao, CHOU Gui-Sheng, ZHANG Huai-Jiang, MA Chun-Sen
    2012, 55(10):  1230-1238. 
    Abstract ( 3614 )   PDF (1408KB) ( 1166 )     
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    In order to understand the influence of Malus sieversii subsp. kirghisorum and M. domestica Golden Delicious on the development and reproduction of the European red mite, Panonychus ulmi (Koch), age-stage two-sex life tables of the experimental population of P. ulmi on the two varieties under the conditions of 23±1℃, 75%±5% relative humidity and a photoperiod of 16L∶8D in the laboratory were constructed. The results showed that there were significant differences in female adult longevity, oviposition period and fecundity between P. ulmi mites reared on M. sieversii subsp. kirghisorum and on M. domestica Golden Delicious (P<0.05). However, no significant difference was found in total developmental duration, total pre-oviposition period and survival rate of immatures between P. ulmi mites on M. sieversii subsp. kirghisorum and on M. domestica Golden Delicious. The total developmental duration of P. ulmi on M. sieversii subsp. kirghisorum and M. domestica Golden Delicious were 12.60 d and 12.54 d for females, and 11.40 d and 11.67 d for males, respectively. The longevity of female P. ulmi mites reared on M. domestica Golden Delicious (13.46 d) was longer than those on M. sieversii subsp. kirghisorum (10.88 d). The oviposition periods of P. ulmi on M. domestica Golden Delicious and M. sieversii subsp. kirghisorum were 10.55 d and 8.3 d, respectively. The number of eggs laid per female was significantly higher on M. domestica Golden Delicious (34.12 eggs/female) than on M. sieversii subsp. kirghisorum (22.48 eggs/female) (P<0.05). The intrinsic rate of increase (r), the net reproductive rate (R0), the mean generation time (T) and the finite rate of increase (λ) of P. ulmi mites reared on M. sieversii subsp. kirghisorum were 0.1354, 11.96, 18.33 and 1.1450, respectively, while those reared on M. domestica Golden Delicious were 0.1489, 17.39, 19.18 and 1.1606, respectively. The results of population dynamic parameters suggest that the population growth of P. ulmi mites reared on M. domestica Golden Delicious is quicker than those reared on M. sieversii subsp. kirghisorum. The results may contribute to further understanding of the population dynamics of P. ulmi occurring on M. sieversii and M. domestica, and provide a theoretical basis to breed mite-resistant apple varieties and to promote comprehensive management for P. ulmi.