Host-parasite relationships of the genus Hyalomma Koch, 1844 (Acari, Ixodidae) and their connection with microevolutionary process

Host-parasite relationships of Hyalomma species of the world fauna are analyzed. The majority of species infests predominately various mammals. Birds and reptiles are used as preferred hosts by several Hyalomma species, and only on certain stage: adults of H. aegyptium parasitize tortoises; immature stages of H. marginatum parasitize birds. It is hypothesized that relationships of H. aegyptium adults (subgenus Hyalomma s. str.) with reptiles are secondarily in origin. Immature stages of H. aegyptium retain the primary wide diapason of hosts, which are various small mammals, birds and reptiles. The life cycle of this species is the three-host type that is considered as a primary type in ixodid ticks. A typical scheme of relationships with their hosts in all well-examined Hyalommina species has following features: the adult stage parasitize large and medium sized mammals, immature stages parasitize small mammals, three-host life cycle. A variety of preferred hosts and types of life cycle is observed in the subgenus Euhyalomma. All species of this subgenus can be arranged into two groups. In the first group, the immature stages infest only small mammals and birds, and the adults parasitize large mammals; this type of host preferences is probably primary host-parasite relationships of Hyalomma. This group includes: H. albiparmatum, H. asiaticum, H. excavatum, H. franchinii, H. impeltatum, H. impressum, H. lusitanicum, H. marginatum, H. nitidum, H. schulzei, and H. truncatum. Hyalomma marginatum and H. schulzei are two-host species; H. excavatum is two- or three-host tick. All the remaining species (except H. albiparmatum, which life cycle is unknown) are three-host ticks. In the second group, the immature stages as well as the adult stage parasitize large mammals. This group includes: H. dromedarii, H. anatolicum, and H. scupense. These species are two- or one-host ticks.     

Larval identification of species and subspecies of the genus Hyalomma (Acari: Ixodidae) from Russia and neighbouring territories

The following species and subspecies of Hyalomma Koch, 1844 are recorded from Russia and neighbouring territories, including those involved in natural foci of tick-borne diseases: H. (Hyalomma) aegyptium (Linnaeus, 1758), H. (Euhyalomma) dromedarii Koch, 1844, H. (Euh.) asiaticum asiaticum Schulze et Schlottke, 1930, H. (Euh.) asiaticum caucasicum Pomerantzev, 1940, H. (Euh.) asiaticum kozlovi Olenev, 1931, H. (Euh.) anatolicum Koch, 1844, H. (Euh.) excavatum Koch, 1844, H. scupense Schulze, 1918, H. (Euh.) marginatum marginatum Koch, 1844, H. (Euh.) marginatum turanicum Pomerantzev, 1946, and H. (Euh.) marginatum rufipes Koch, 1844. The geographic distribution and host-parasite relationships of each taxon are discussed. Species characters of the larval stage, that can be distinguished using light microscopy, are found to be very few. These characters include shape of scutum, shape and rate of hypostome denticulation, shape and rate of the development of spurs on coxae I to III. Measurements of some morphological structure and their rations show statistically significant differences between some closely related species, even if qualitative discriminating characters are unknown. Only measurements and their ratios can be used for the discrimination of larval H. anatolicum from larval H. excavatum, because qualitative discriminating features have not been found for these species. The complex structure of the subspecific morhological differentiation of all parasitic stages in the polymorphic species H. asiaticum and H. marginatum is revealed. This structure probably reflects some peculiarities of the microevolutionary processes. Identification key for the larval stage of seven Hyalomma species is provided.     

Ornithodoros (Alectorobius) spheniscus n. sp. [Acarina: Ixodoidea: Argasidae: Ornithodoros (Alectorobius) capensis group], a tick parasite of the Humboldt penguin in Peru

Ornithodoros (Alectorobius) spheniscus n. sp., described from wild-caught and laboratory-reared females, males, nymphs, and larvae parasitizing the Humboldt Penguin, Spheniscus humboldti Meyen, is the fifth species of the Ornithodoros (Alectorobius) capensis group to be recognized in the Neotropical Region. A related Peruvian species, Ornithodoros (Alectorobius) amblus Chamberlin, also parasitizes S. humboldti but is recorded from a wider range of marine birds breeding on the Pacific coast and offshore islands, where the birds congregate to feed on the rich fish fauna usually produced by the Humboldt current. Differential criteria are provided for the new species, O. (A.) amblus, and Ornithodoros (Alectorobius) yunkeri Keirans, Clifford, and Hoogstraal of the Galapagos. These 3 members of the O. (A.) capensis group parasitize marine birds associated with the Humboldt current in western South America and the Galapagos. Persons visiting Humboldt Penguin breeding sites in caves and on barren coastal ledges are eagerly attacked by nymphal and adult O. (A.) spheniscus and suffer afterward from pruritus and slowly-healing blisters. The O. (A.) spheniscus life cycle required 128 to 193 days in the laboratory and, as typical of bird-parasitizing members of the subgenus Alectorobius, the first nymphal instar did not feed.     

Detection of Dichelobacter nodosus in wild ungulates (Capra ibex ibex and Ovis aries musimon) and domestic sheep suffering from foot rot using a two-step polymerase chain reaction

Severe keratinous hoof afflictions have been recorded in ibex (Capra ibex ibex) since 1995 and more recently in mouflon (Ovis aries musimon) in Switzerland. Based on clinical observations and comparison with diseases known to affect domestic ungulates, it was hypothesized these wild ungulates were affected by foot rot associated with infection with Dichelobacter nodosus. Dichelobacter nodosus has been shown to be the essential pathogen for initiation and establishment of foot rot, a highly contagious foot disease of sheep and goats. Because these bacteria could not be cultivated from affected ibex, we developed a nested polymerase chain reaction that allowed detection of D. nodosus without culture. Using this assay, we were able to diagnose D. nodosus infections of ibex, mouflon, and domestic sheep in natural outbreaks. From these results we conclude that D. nodosus plays an etiological role in foot rot not only in domestic but also in wild Caprinae.     

Distribution and ecology of ticks (Acari: Ixodidae) infesting livestock in Tunisia: an overview of eighth years field collections

Ticks (Ixodidae) play a significant role as vectors of pathogens of domestic animals in Tunisia. The major losses caused by ticks are related to transmission of protozoan parasites. These include agents of tropical theileriosis and babesiosis in ruminants. Since 1991, we conducted research studies on tick population of livestock in Tunisia. This overview reports a synthesis on tick distribution, their biology and their role as vectors of pathogens in domestic animals, particularly cattle. During the whole period of the study about 15,000 tick specimens were collected from different zones of the country. A total of 14 species were identified. Hyalomma detritum detritum was the most abundant and important (vector of Theileria annulata) species infesting cattle. Hyalomma dromedarii and Hyalomma impeltatum were collected on domestic ruminants in the arid and desertic zones. Hyalomma marginatum marginatum and Hyalomma anatolicum excavatum were widespread and found on livestock hosts. Ixodes ricinus, vector of Babesia divergens and Borrelia burgdorferi sl, colonises mainly the humid zone. Boophilus annulatus and Rhipicephalus bursa infesting cattle, sheep and goats were found in the sub-humid and semi-arid zones. Haemaphysalis sulcata and Hae. punctata were collected in humid and sub-humid zones on cattle and sheep. Rhipicephalus turanicus were collected in different regions, on different animal species. Rhipicephalus sanguineus, tick of dogs, were often collected on livestock. Only few specimens of Hyalomma marginatum rufipes and Hyalomma franchinii were collected.     

Field and vegetable crops as hosts of larval western spotted cucumber beetle (Coleoptera: Chrysomelidae)

The western spotted cucumber beetle, Diabrotica undecimpunctata undecimpunctata Mannerheim, is an important pest of melons (Cucurmis melo L.) in northern California. Recent observations indicate that adults are using alfalfa (Medicago sativa L.) as a feeding host and larvae may be developing on the roots. Greenhouse studies were conducted during the winters of 2009 and 2010 in which larval development was compared on the roots of six field and vegetable crops commonly grown in the southern Sacramento Valley. The growth parameters used to evaluate the hosts were larval and pupal head capsule width, body width, and body length as well as total survival percentage and survival percentage to the third instar. According to larval growth and survivorship in 2009, maize (Zea mays L.) was the best host, followed by alfalfa and tomato (Solanum lycopersicum L.) that were roughly equivalent to one another. Melon was a slightly weaker host than alfalfa and tomato; sunflower (Helianthus annuus L.) was a completely incompatible host and thus dropped from the 2010 study. In 2010, melon was the weakest host for larval development. Maize was the superior host again followed closely by alfalfa that performed slightly better than tomato and lima bean (Phaseolus lunatus L.). Data suggest that larval western spotted cucumber beetles may primarily develop outside of melon fields and the adults emigrate to melons.     

The distribution of Hyalomma spp. ticks from domestic ruminants in Iran

The distribution of Hyalomma species on domestic animals was studied in four zoogeoghraphical zones. Nine hundred and ninety-two Hyalomma ticks were collected from sheep, goats, cattle and camels. A total of seven tick species consisting of: Hyalomma anatolicum Koch, 1844 (28.93%), H. excavatum Koch, 1844 (12.5%), H. asiaticum Schulze & Schlottke, 1930 (13.5%), H. marginatum Koch, 1844 (27.01%), H. detritum Schulze, 1919 (9.67%), H. schulzei Olenev, 1931 (4.03%) and H. dromedary Koch, 1844 (4.33%) were recorded. The results indicated that H. anatolicum, H. asiaticum, H. marginatum and H. detritum were present in all zones whereas H. excavatum was absent in Zone I. The result also showed that H. dromedari was presented in Zones III and IV whereas H. schulzei was present in Zones II and III. During this study, the tick species collected are candidates for investigation as vectors of pathogenic organisms in Iran.     

The genus Hyalomma. XI. Redescription of all parasitic stages of H. (Euhyalomma) asiaticum (Acari: Ixodidae) and notes on its biology

The tick Hyalomma (Euhyalomma) asiaticum Schulze & Schlottke is provisionally considered to belong to the H. (E.) asiaticum group of closely related species. Males of H. asiaticum can be distinguished from those of other species of the group by their long and very deep cervical grooves, long, narrow, straight adanal plates, long dorsal prolongation of the spiracular plates, dorsal posterior margin of the basis capituli deeply concave and angular, and unbroken ivory-coloured strip on the dorsal aspect of the leg segments. Females of H. asiaticum can be distinguished from those of other species of the H. asiaticum group by their very deep cervical grooves, narrowly U-shaped genital aperture, with bulging preatrial fold. Larger domestic and wild ungulates are the principal hosts of the adults, while nymphs and larvae parasitize mainly rodents, leporids and hedgehogs. Hyalomma asiaticum is widely distributed in Asia, from Syria in the West to eastern China in the East. Here all the parasitic stages of H. asiaticum are illustrated and redescribed. Data on its disease relationships are also provided.     

Metazoan parasites on the gills of the skipjack tuna Katsuwonus pelamis (Osteichthyes: Scombridae) from the Alboran Sea (western Mediterranean Sea)

The gills of 31 skipjack tuna Katsuwonus pelamis (L.) caught in the Alboran Sea (western Mediterranean Sea) were examined for metazoan parasites, and the gills of 4 specimens from the Balearic Sea (also western Mediterranean Sea) were analysed for comparative purposes. Nine -species of parasites were found, including 8 didymozoid trematodes (Atalostrophion cf. bio-varium, Didymocylindrus filiformis, Didymocylindrus simplex, Didymocystis reniformis, Didymoproblema fusiforme, Didymozoon longicolle, Koellikeria sp. and Lobatozoum multisacculatum) and 1 caligid copepod (Caligus bonito). Koellikeria sp. and L. multisacculatum were not recorded in the Balearic Sea. Most of the parasites (79.2% of all specimens) were didymozoids. Didymozoon longicolle was the dominant species; A. cf. biovarium, D. simplex, D. fusiforme and L. multisacculatum are reported from the Mediterranean Sea for the first time. No correlation was found between the intensity of infection of any parasite species and host size or sex. Most of the parasites, particularly didymozoids, showed a high site-specificity. Significant differences were found between the parasite assemblages of K. pelamis from the Alboran Sea and from the Atlantic Ocean. D. fusiforme, D. longicolle and L. multisacculatum are suggested as potential tags to follow skipjack tuna migrations between the Atlantic Ocean and Mediterranean Sea.     

The seasonal occurrence of ticks (Acari: Ixodidae) on sheep and in the field in the Judean area of Israel

A 2 year survey of ixodid ticks in the Judean area of Israel between 1983 and 1985 showed that sheep were parasitized by the following species:Rhipicephalus sanguineus group (30.6%),Rhipicephalus bursa (25.3%),Haemaphysalis cretica (20.5%),Haemaphysalis otophila (20.1%) andHyalomma anatolicum excavatum (3.1%). Three other species,Hyalomma marginatum rufipes, Boophilus annulatus andAmblyomma lepidum were also present but in very low numbers. A total of 50 816 specimens (78.1% adult ticks) were collected from the sheep. In the field, a total of 4780 ticks (8.7% adults) were collected and 64.3% consisted ofR. bursa. Other species found wereH. cretica (14.5%),H. a. excavatum (13.7%),H. otophila (3.4%),R. sanguineus group (1.9%) and 1.7% wasIxodes eldaricus. The number of ticks collected varied from year to year and seasonal activity was observed in all species except forH. a. excavatum. In all species, males outnumbered females, with the exception ofH. cretica. The seasonal occurrence, tick ecology and disease relationship is discussed for each species.     

Transmission efficiency of Xylella fastidiosa by sharpshooters (Hemiptera: Cicadellidae) in coffee and citrus

Xylella fastidiosa (Wells, Raju, Hung, Weisburg, Mandelco-Paul, and Brenner) is a bacterial pathogen transmitted by several sharpshooters in two tribes of Cicadellinae (Proconiini and Cicadellini). Here, we compared the transmission efficiency of X. fastidiosa in coffee (Coffea arabica L.) and citrus [Citrus sinensis (L.) Osbeck] by Cicadellini [Bucephalogonia xanthophis (Berg) and Dilobopterus costalimai Young] and Proconiini [Homalodisca ignorata Melichar and Oncometopia facialis (Signoret)] sharpshooters that occur in both crops. At different seasons, healthy adults of each species were submitted to a 48-h acquisition access period on citrus or coffee source plants infected with X. fastidiosa isolates that cause Citrus variegated chlorosis (CVC) and Coffee leaf scorch (CLS), respectively, and then confined on healthy seedlings of the corresponding host plant for a 48-h inoculation access period. No significant effect of inoculation season was observed when comparing infection rates of citrus or coffee plants inoculated by vectors at different times of the year. In citrus, the transmission rate by single insects was significantly higher for H. ignorata (30%) in relation to B. xanthophis (5%) and O. facialis (1.1%), but there was no difference among vector species in coffee, whose transmission rates ranged from 1.2 to 7.2%. Comparing host plants, H. ignorata was more effective in transmitting X. fastidiosa to citrus (30%) in relation to coffee (2.2%), whereas the other vectors transmitted the bacterium to both hosts with similar efficiencies. Despite these variations, vector efficiency in coffee and citrus is lower than that reported in other hosts.     

The life cycle of Hyalomma rufipes (Acari: Ixodidae) under laboratory conditions

Biological characteristics of Hyalomma rufipes parasitising on rabbits and sheep were compared under laboratory conditions in Gansu, China. Mature ticks could parasitize both rabbits and sheep, while immature ticks only fed on rabbits successfully. Adults sucked blood on sheep significantly longer than on rabbits (16 and 13 days, respectively). Other adult parasite characteristics fed on the two hosts were similar, including the weight of engorged adult, female daily oviposition, and the weight and amount of the egg mass laid. Those indicated that this tick species showed little host specificity between sheep and rabbits during its adult stage. In total, the life cycle of H. rufipes was completed in an average period of 179 days. The average developmental periods were 59 days for egg incubation, 3 and 21 days for immature tick prefeeding and feeding, 2, 12 and 40 days for adult prefeeding, female preoviposition and oviposition. The longer female fed for engorgement, the shorter preoviposition period of engorged female needed, although when the feeding period was less than 15 days, this relationship was not obvious. The results confirmed the correlation between the weight of the engorged female and the number of eggs laid (r = 0.909). The reproductive efficiency index (REI) and reproductive fitness index (RFI) in females was 10.63 and 7.22, respectively. Engorged nymphs moulting to females were significantly heavier (27.6 ± 0.89 mg) than those moulting to males (22.3 ± 0.52 mg). Males outnumbered females by 1.4:1.     

Effects of Pieris host species on life history parameters in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

Host specificity and host selection by insect parasitoids are hypothesized to be correlated with suitability of the hosts for parasitoid development. The present study investigates the correlation between host suitability and earlier studied host-finding behaviour of two closely related braconid larval parasitoid species, the generalist Cotesia glomerata (L.) and the specialist C. rubecula (Marshall) (Hymenoptera: Braconidae). We compared the capability of both parasitoid species to parasitize and develop in three Pieris host species, i.e. P. brassicae (L.), P. rapae (L.) and P. napi (L.) (Lepidoptera: Pieridae). In laboratory experiments, we measured the effect of host species on fitness parameters such as survival, development, sex ratio and size of parasitoid progeny. The results show that C. glomerata is capable of developing in the three host species, with significant differences in parasitoid survival, clutch size and adult weight among Pieris species. The host range for development was more restricted for C. rubecula. Although C. rubecula is physiologically able to develop in P. brassicae larvae, parasitoid fitness is negatively affected by this host species, compared to its most regular host, P. rapae. A comparison of the present data on host suitability with earlier studies on host-searching behaviour suggests that the host-foraging behaviour of both parasitoid species not only leads to selection of the most suitable host species for parasitoid development, but also plays a significant role in shaping parasitoid host range.     

Towards a diagnostic view of Hyalomma (Hyalomma) aegyptium (Acari,Ixodidae)

The diagnostic characters of larval, nymphal and adult Hyalomma aegyptium (L., 1758) based on specimens from the territory of most part of the area are given. In the diagnoses of immature stages, was used the characters, which were formerly tested by the author for diagnostics of other Hyalomma species occurring in the former USSR. Commonly used characters and those which were revealed by the author as useful for the majority of Euhyalomma Filippova, 1984 and Hyalommina Schulze, 1919 species, have been used in the diagnoses of male and female. Differential diagnosis of Hyalomma aegyptium. Female: genital orifice as wide arch with straight posterior margin (fig. 2, 5); vestibular part of vagina funnel-like, greatly swollen (fig. 2, 5); setae of alloscutum stick-like, tapering in apical one (figs 2, 3, 4); second segment of palps with proximal narrowing (figs 3, 1, 2); spurs of coxae I widely separated, triangular, wide, subequal in size (fig. 3, 5). Male: any grooves of conscutum absent, except short and pit-like cervical ones (fig. 4); punctation sparce and impressive (fig. 4); adenal shields short and wide, without inner branch, posterior part widened, anteromedian margin straight (fig. 5, 4); spurs of coxae I widely separated, triangular, wide, subequal in size (fig. 6, 5). Nymph: posteromedian setae of alloscutum stick-like and, as a rule, with indented apices (fig. 7, 2); spurs of coxae I large, median spur as equilateral triangle and shorter than lateral one (fig. 7, 8); spurs of coxae II-IV well developed, with acute apices (fig. 7, 8). Larva: posterior part of scutum (behind the eyes) heavy elongated, its apex straight, postero-lateral incisions weakly developed (fig. 8, 1); spurs of coxae I as equilateral triangle in shape and with rounded apices, spurs of coxae II-III very large (fig. 8, 5).     

Evaluation of host range and larval feeding impact of Chrysolina aurichalcea asclepiadis (Villa): considerations for biological control of Vincetoxicum in North America

A biological control program has been initiated against European swallow-worts Vincetoxicum nigrum (L.) Moench. and V. rossicum (Kleopow) Barbar., which are invasive in North America. A population of the leaf beetle Chrysolina aurichalcea asclepiadis (Villa) originating from the western Alps has been under evaluation as a part of this program. The preliminary host range of C. a. asclepiadis was determined among 37 potential host plants. In addition, a prerelease impact study was conducted to determine the effect of larval feeding on the performance of V. nigrum. Under no-choice conditions beetle larvae completed development on nine plant species within the genera Artemisia and Tanacetum (Asteraceae) and Asclepias and Vincetoxicum (Apocynaceae). The host range of adults is broader than larvae (13 plant species within five genera received sustained feeding). Three of the six nontarget species supporting larval development are native to North America, however in separate oviposition tests, female beetles failed to produce eggs when confined to these hosts. In multiple-choice tests, neither larvae nor adults preferred Vincetoxicum spp. to nontarget species. Larval damage by C. a. asclepiadis at densities at and above five larvae per plant substantially reduced growth, biomass, and delayed reproduction of V. nigrum. However, this population of C. a. asclepiadis is polyphagous and unsuitable for biological control of Vincetoxicum because of potential risk of attack to Asclepias tuberosa L. and native North American Asteraceae, particularly Artemisia.     

Parasitism of three attelabid weevils by Bracon (Uncobracon) apoderi Watanabe (Hymenoptera: Braconidae) in Kyushu,Japan

The larval parasitoid Bracon (Uncobracon) apoderi Watanabe (Hymenoptera: Braconidae) is reported from Kyushu for the first time. Two host weevils, Phymatapoderus latipennis and Paroplapoderus pardalis (Coleoptera: Attelabidae), are newly recorded. Bracon apoderi parasitized three attelabid species, including Apoderus balteatus, but did not parasitize Cycnotrachelus roelofsi even though it is a dominant attelabid species in the study site. In addition, B. apoderi mainly parasitized suspended cradles rather than cut‐off cradles.     

Co-distribution pattern of a haemogregarine Hemolivia mauritanica (Apicomplexa: Haemogregarinidae) and its vector Hyalomma aegyptium (Metastigmata: Ixodidae)

Hyalomma aegyptium ticks were collected from tortoises, Testudo graeca, at localities in northern Africa, the Balkans, and the Near and Middle East. The intensity of infestation ranged from 1-37 ticks per tortoise. The sex ratio of feeding ticks was male-biased in all tested populations. Larger tortoises carried more ticks than did the smaller tortoises. The juveniles were either not infested, or carried only a poor tick load. Hyalomma aegyptium was absent in the western Souss Valley and Ourika Valley in Morocco, the Cyrenaica Peninsula in Libya, Jordan, and the Antilebanon Mountains in Syria. Hemolivia mauritanica, a heteroxenous apicomplexan cycling between T. graeca and H. aegyptium, was confirmed in Algeria, Romania, Turkey, Syria, Lebanon, and Iran. Its prevalence ranged from 84% in Romania (n = 45), 82% in eastern Turkey (n = 28), and 82% in the area of northwestern Syria with adjacent Turkish borderland (n = 90), to 38% in Lebanon (n = 8) and in only 1 of 16 sampled tortoises in Algeria. The intensity of parasitemia in the studied areas ranged from 0.01% up to 28.17%. The percentage of Hemolivia-infected erythrocytes was significantly higher in adults. All tortoises from Hyalomma-free areas were Hemolivia-negative. Remarkably, all 29 T. graeca from Jabal Duruz (southwestern Syria) and 36 T. graeca from the area north of Middle Atlas (Morocco) were Hemolivia-negative, despite the fact that ticks parasitized all adult tortoises in these localities. Identical host preferences of H. aegyptium and H. mauritanica suggest the occurrence of co-evolution within the Testudo-Hyalomma-Hemolivia host-parasite complex.     

缘叶舌蜂与其天敌窄头褶翅蜂的生物学特性 及其与环境变量的相关性

【目的】记录亚热带地区独栖性膜翅目昆虫的生物学特性,研究乔木树种多样性对缘叶舌蜂Hylaeus perforata及其天敌窄头褶翅蜂Gasteruption corniculigerum的影响。【方法】2015年8月至2018年9月,在江西省德兴市新岗山镇的生物多样性与生态系统服务功能实验样地人工林设置了标准化的人工巢管收集缘叶舌蜂与窄头褶翅蜂,观察其生物学特性,分析缘叶舌蜂和天敌窄头褶翅蜂多度以及窄头褶翅蜂寄生率与乔木树种丰富度、海拔、坡度、北向指数和东向指数5个环境变量的相关性。【结果】缘叶舌蜂一年多代,每年5-9月筑巢,4-8月羽化活动;使用树脂筑巢;平均每次建造巢室3.75±2.24个;巢的平均直径4.73±1.31 mm;以末龄幼虫越冬;后代性比显著偏向雌性。窄头褶翅蜂营盗寄生;每年仅在7月寄生缘叶舌蜂,后代会消耗掉寄主巢内所有蜂粮;9月羽化活动;羽化时间远晚于同期的寄主缘叶舌蜂。缘叶舌蜂和窄头褶翅蜂的多度与树种丰富度呈正相关,而其他4个环境变量对缘叶舌蜂和窄头褶翅蜂的多度没有显著影响。【结论】缘叶舌蜂是典型的独栖性蜜蜂,每次筑巢会产下多个后代,在春夏及初秋活动;窄头褶翅蜂在夏季寄生缘叶舌蜂,秋季羽化活动,发育时间远长于寄主,其他盗寄生类群一般寄生单个寄主的巢室,而窄头褶翅蜂会将寄主的所有巢室破坏并消耗掉寄主储存的所有蜂粮;树种多样性更高的环境更有利于高营养级的物种生存。