Host location,survival and fecundity of the Oriental rat flea Xenopsylla cheopis (Siphonaptera: Pulicidae) in relation to black rat Rattus rattus (Rodentia: Muridae) host age and sex

Host choice and fecundity are two factors that may contribute to the variation in flea counts observed when assessing the potential risk of flea-borne transmission of pathogens from rodents to humans. Using the black rat, Rattus rattus Linnaeus, as host the effects of age and sex on host choice and fecundity of the Oriental rat flea, Xenopsylla cheopis Rothschild, were examined experimentally at 25 degrees C and 80% rh. During the first two days of emergence from cocoons, female fleas dominated the sex ratio by 4:1 but from the third day onwards this switched to a male-dominated sex ratio of 4:1. The sex of the flea did not influence their host-seeking behaviour. Newly emerged fleas of both sexes were not influenced by the rat's presence and at seven days old both sexes demonstrated similar levels of attraction toward the rat host. The sex of the rat did not affect flea host-seeking behaviour. There was a 50-70% decline in the initial number of adult fleas during the first week after their release onto a rat host, and this decline was greatest on juvenile rats. Flea fecundity was also significantly lower on juvenile rat hosts but no differences due to the sex of the rat were observed. This experimental study supports the hypothesis that differences in flea count due to host sex, reported in field surveys, result from sexual differences in host behaviour and not from discriminatory host-seeking behaviour by X. cheopis. Differences in flea count due to host age may be affected by differences in X. cheopis fecundity, which may itself be mediated by host behaviour such as grooming.     

The longevity of Leptopsylla segnis fleas (Siphonaptera: Leptopsyllidae)

In experiments, the mean life duration of fleas Leptopsylla segnis on white mice (abundance of fleas within natural limits, up to 10 fleas per mouse) was 22.7 days in females and 18.8 day in males. Maximum life duration was 51 and 37 days respectively. In cases, when the initial numbers of fleas were 20 and 28-34 fleas, the duration of life was decreased. The maximum limit decreased greater than the mean duration of life. A survival dynamics of fleas depended upon the flea number. It was found out, that in cases of high abundance of fleas in the beginning of experiments, the mortality rate of males was lower than in females. During the stay on a host the fleas lost gradually an ability to endure a starvation. Possible mechanisms of the regulation of flea abundance are discussed.     

Development rates of two Xenopsylla flea species in relation to air temperature and humidity

The rate of development of immature fleas, Xenopsylla conformis Wagner and Xenopsylla ramesis Rothschild (Siphonaptera: Xenopsyllidae) was studied in the laboratory at 25 degrees C and 28 degrees C with 40, 55, 75 and 92% relative humidity (RH). These fleas are separately associated with the host jird Meriones crassus Sundevall in different microhabitats of the Ramon erosion cirque, Negev Highlands, Israel. This study of basic climatic factors in relation to flea bionomics provides the basis for ecological investigations to interpret reasons for paratopic local distributions of these two species of congeneric fleas on the same host. Both air temperature and RH were positively correlated with duration of egg and larval stages in both species. Change of humidity between egg and larval environments did not affect duration of larval development at any temperature. At each temperature and RH, the eggs and larvae of X. ramesis did not differ between males and females in the duration of their development, whereas female eggs and larvae of X. conformis usually developed significantly faster than those of males. For both species, male pupae developed slower than female pupae at the same air temperature and RH. Air temperature, but not RH, affected the duration of pupal development. At each humidity, duration of the pupal stage was significantly longer at 25 degrees C than at 28 degrees C: 15.3+/-1.7 vs. 11.7+/-1.2 days in X. conformis; 14.1+/-2.0 vs. 11.5+/-1.7 days in X. ramesis, with a significantly shorter pupal period of the latter species at 25 degrees C. These limited interspecific bionomic contrasts in relation to basic climatic factors appear insufficient to explain the differential habitat distributions of X. conformis and X. ramesis.     

Experimental transmission of murine typhus by Xenopsylla cheopis flea bites

Transmission of Rickettsia typhi to rats by the bites of Xenopsylla cheopis (Rothschild) fleas was investigated. Procedures rigorously excluded the possibility of contamination of the host skin by flea faeces. Fleas with R. typhi infection (21-25 days post-infection) which fed through bolting cloth (45 min exposure to ten fleas) transmitted rickettsiae with a success rate of 20%. Infective fleas allowed free access to their host for 8 h (10-15 fleas/rat) gave transmission rates of 45-68%. They were also capable of inoculating R. typhi through a membrane of rat skin on a feeder. Only fleas which had been infected for 21 days or longer transmitted R. typhi orally. Oral transmission appeared to be the result of regurgitation of rickettsiae present in the foregut lumen rather than through salivary secretions.     

北方几种蚤类侵袭寄主的研究和斧形盖蚤叮人试验

本文研究了二齿新蚤和方形黄鼠蚤松江亚种对寄主的侵袭活动,包括活寄主与死寄主被侵袭情况比较,寄主体表蚤数与环境温度的关系,以及与蚤同寄主接触时间的关系。还研究了达乌尔黄鼠和达乌尔鼠兔寄生蚤在一日不同时间里的数量变化。并观察了斧形盖蚤叮人情况。     

Life expectancy of Ctenophthalmus wladimiri Is.-Gurv., 1948 (Siphonaptera, Ctenophthalmidae) under laboratory conditions]

Laboratory studies have shown that at 75% humidity fleas, Ct. wladimiri, are less viable than at 100% humidity. At 100% humidity at a temperature from 0 to 10 degrees and without feeding newly born imagos survived up to 363 days, at a higher temperature from 18 to 30 degrees-from 44 to 163 days. At a temperature from 7 to 10 degrees imagos, which fed one time, lived up to 355 days and at 0 to 2 degrees-up to 287 days. Maximum life duration (1133 days) for periodically feeding fleas was at 100% humidity at a temperature from 7 to 10 degrees. Under such conditions 50% of fleas survived up to 114 days. Hungry but previously "prepared" for winter fleas lived at a temperature from 0 to 2 degrees not more than 376 days.     

Effect of temperature and relative humidity on the development and fecundity of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae)

The spotted stemborer, Chilo partellus (Swinhoe) (Lepidoptera: Crambidae), is one of the most important insect pests attacking maize and sorghum in Ethiopia. Recent studies have indicated that the pest is spreading to new locations where it was not reported before. In the current study, laboratory investigations were carried out to determine the combined effect of different levels of relative humidity and temperature regimes on the development and fecundity of C. partellus, as these physical factors are known to play an important role in the life cycle of insects and adaptability to local climate. Developmental time, longevity, potential fecundity and realized fecundity of C. partellus were measured under controlled conditions. Three temperature regimes (22°C, 26°C and 30°C) and three relative humidity levels (40%, 60% and 80%) were tested. It was found that temperature, relative humidity (RH) and their interaction significantly affected the developmental time, adult longevity, potential fecundity and realized fecundity of the pest. Developmental time was inversely related to temperature. Mean duration of C. partellus life cycle was 70.2 days at 22°C and 80% RH, whereas it took only 26.5 days to complete its life cycle at 30°C and 40% RH. Male and female longevity were similar in most cases. The adult life span ranged between 6.9-11.1 days at 22°C and 3.1-7.2 days at 30°C for different levels of relative humidity. The most suitable conditions for C. partellus development and fecundity were 26-30°C temperatures regimes and 60-80% RH levels.     

Effects of fleas on nest success of Arctic barnacle geese: experimentally testing the mechanism

Parasites have detrimental effects on their hosts’ fitness. Therefore, behavioural adaptations have evolved to avoid parasites or, when an individual is already in contact with a parasite, prevent or minimize infections. Such anti‐parasite behaviours can be very effective, but can also be costly for the host. Specifically, ectoparasites can elicit strong host anti‐parasite behaviours and interactions between fleas (Siphonaptera) and their hosts are one of the best studied. In altricial bird species, nest fleas can negatively affect both parent and offspring fitness components. However, knowledge on the effects of fleas on precocial bird species is scarce. Research on geese in the Canadian Arctic indicated that fleas have a negative impact on reproductive success. One possible hypothesis is that fleas may affect female incubation behaviour. Breeding females with many fleas in their nest may increase the frequency and/or duration of incubation breaks and could even totally desert their nest. The aim of our study was to 1) determine if a similar negative relationship existed between flea abundance and reproductive success in our study colony of Arctic breeding barnacle geese Branta leucopsis and 2) experimentally quantify if such effects could be explained by a negative effect of nest fleas on female behaviour. We compared host anti‐parasite and incubation behaviour between experimentally flea‐reduced and control nests using wildlife cameras and temperature loggers. We found that flea abundance was negatively associated with hatching success. We found little experimental support, however, for changes in behaviour of the breeding female as a possible mechanism to explain this effect.     

The Ectoparasites of the European Badger, Meles meles, and the Behavior of the Host-Specific Flea, Paraceras melis

The badger-specific flea, Paraceras melis, jumps repeatedly when separated from its host Meles meles; thereafter fleas settled into sheltered positions. After separation from badgers, some 42% of fleas (n = 63) voided their gut contents; this was associated with a significant increase in mean jumping distance. The maximum longevity of fleas away from the host was 89 days, with 50% mortality at around 35 days. Badger lice, Trichodectes melis, survived for up to 3 days postcapture. We conclude that the badger's habit of frequently swapping dens with a mean period of return of 6 days is unlikely to bring about significant mortality of adult fleas but may effectively eradicate lice. Fleas abandoned in bedding in a simulated badger sett were mobile, being drawn toward light and moving upward. This response would draw the fleas to the den entrance, which may be a suitable site to intercept returning badgers. The fleas responded to stimuli which might signal the proximity of the host: they jumped toward sources of carbon dioxide and of carbon dioxide in air current directed at the flea. The strongest response was seen when a mixture of stimuli consisting of carbon dioxide, a dark circle of card, and movement were tested; the majority of fleas jumped toward the mixed stimulus. Finally, fleas separated from the host responded to exhaled air by running and jumping; this is in marked contrast to their response to those stimuli when on the host, when fleas run downward and very rarely jump. These contrasting observations find adaptive explanation in the two contexts.     

Notes on the life-history of the rabbit flea Caenopsylla laptevi ibera Beaucournu & Marquez, 1987 (Siphonaptera: Ceratophyllidae) in eastern Spain

The rabbit flea Caenopsylla laptevi ibera occurs in arid environments of central and eastern Spain. Although the fleas breed during the coolest, most humid part of the year, the larvae survive and grow in sand at only 50-60% relative humidity. At 22 degrees C and 80% relative humidity eggs hatch in six days and the cocoon stage is reached 10-11 days after hatching. Female fleas emerge from pupation at about 17 days after cocoon spinning; males emerge a little later at a mean of 20 days. Adult fleas are mainly found on the host Oryctolagus cuniculus. Measurements of burrow microclimate confirmed that in south-eastern Spain burrow humidity was adequate for the development of C. I. ibera larvae over most of the year. However, breeding may be restricted for at least part of the year, as the larvae of C. I. ibera apparently cannot complete development at 25 degrees C or above. In the laboratory, fleas can enter a prolonged quiescent period while in the cocoon. This is possibly a facultative, pre-pupal diapause and the likely mechanism that accounts for the disappearance of adult fleas from the field by spring and their reappearance each autumn.     

An attempt to protect cats against infestation with Ctenocephalides felis felis using gut membrane antigens as a vaccine

Cats (n = 5) were vaccinated with membrane antigens extracted from the gut of unfed fleas (Ctenocephalides felis felis) together with Quil A and RIBI as adjuvants. Five unvaccinated cats were retained as controls. All the cats were infested on 6 separate occasions with fleas (46–250 per challenge). Protection was assessed from the number of fleas retrieved and the fecundity of the female fleas, measured as the number of developed oocytes contained in the reproductive tract. Cats injected with gut membrane antigens had significantly elevated levels of anti-flea antibodies in their sera, but they were neither protected significantly against infestation with fleas nor was the apparent fecundity of fleas which had fed on vaccinated cats decreased. The possible reason why gut membrane antigens failed to protect cats against fleas are discussed.     

二齿新蚤和方形黄鼠蚤松江亚种侵袭与离开宿主习性的实验研究

本文对二齿新蚤和方形黄鼠蚤松江亚种侵袭与离开宿主的习性进行了实验研究,结果：(1)蚤攻击宿主距离平均在2cm以内,最大攻击距离不超过10cm。在有效侵袭范围内,蚤与宿主距离近时较距离远时吸血蚤数增多。(2)蚤对宿主的侵袭程度雌蚤大于雄蚤,繁殖蚤大于新羽化蚤,二齿新蚤大于方形黄鼠蚤松江亚种,对小白鼠大于对达乌尔黄鼠。(3)宿主死后一定时间内,仍有部份蚤侵袭其尸体。 死亡时间越长侵袭蚤数越少,呈logY=a-blogX型曲线。(4)蚤离开宿主时间呈偏态分布,温度越高偏态分布越明显。蚤离开死鼠和离开活鼠所需时间很接近。 二齿新蚤和方形黄鼠蚤松江亚种离开宿主时间几乎相同。二种蚤离开宿主平均时间与环境温度呈负相关,且呈曲线关系。     

Adult feeding and lifetime reproductive success in the parasitoid Aphytis melinus

The diet of adult females of the parasitoid Aphytis melinus DeBach (Hymenoptera: Aphelinidae) includes host insects and sugar-rich foods such as nectar and honeydew. We compared the contributions of host feeding to longevity and fecundity in A. melinus females in the presence and in the absence of honey meals. First, we assessed the longevity of females that were not allowed to oviposit. While the longevity of females fed honey was significantly increased by host feeding (median ages were 30.5 days for host-fed females and 17 days for females not allowed to host feed), the lifespan of parasitoids not fed honey did not exceed 3 days for any individual and there was no effect of host feeding on longevity in this group. In the second set of experiments, we assessed the fecundity and longevity of females allowed to oviposit. We conducted two experiments, one in which honey was continuously available, and one in which honey was not available. In both experiments, daily observations were made of females that were either allowed to host feed or manually prevented from host feeding. In the presence of honey, host feeding significantly increased both fecundity and longevity, and in the absence of honey, parasitoids died within 2 days and host feeding had no significant effect on either fecundity or longevity. The lifetime fecundity of females fed honey but not hosts exceeded the initial egg complement by 60% on average. Approximately one host per day was used for host feeding whether honey was supplied or not, and each host-feeding meal contributed approximately 3.9 eggs to the lifetime fecundity of honey-fed females. In the last experiment, we compared the rate of egg resorption over a 36-h period in A. melinus females that were deprived of hosts and either fed honey or starved. While no egg resorption was detected in honey-fed females over this time period, starved females resorbed approximately 9 eggs. Thus, the availability of a sugar-rich food interacts strongly with host feeding in influencing longevity and fecundity and has a strong direct effect on egg resorption.     

温度对三叶草彩斑蚜种群参数的影响

为了探讨温度对三叶草彩斑蚜种群增长的影响,在实验室研究了15～35℃范围内共9个温度下三叶草彩斑蚜的发育、繁殖和生命表.结果表明:三叶草彩斑蚜种群在35℃下不能存活.若虫的发育历期随着温度的上升而显著缩短,在15 ～ 32℃内为18.33～4.02 d,存活率在40.0％～83.6％,25℃下若虫的存活率最高、32℃下存活率最低.成蚜的平均寿命在10.64 ～ 20.87 d,23℃下寿命最长、32℃下最短.产蚜高峰期随温度的上升而提前,除15℃为15 d以外,其余温度下均在3～6d.平均繁殖力和单头最高繁殖力以25℃下最高,分别为82.0和149.0头.平均世代周期随着温度的上升从15℃的31.17 d逐渐缩短到32℃的10.17 d.净增殖率在25℃下最大(68.62),32℃下最小(13.96).内禀增长率在0.10 ～0.30 d-1,其中28 ℃下最高、15℃下最低.若虫的发育起点温度和有效积温分别为9.35℃和97.83 d·℃.繁殖力、净增殖率和内禀增长率与温度的关系均可用一元二次方程描述.     

Level of parasitic contact in colonies of the little suslik in the semideserts of western Kazakhstan

Field experiments were conducted during the settling of young animals of Citellus pygmaeus in four areas 4 to 13.6 hectares in size. From 3 to 6 animals were caught in each area and labeled by administering radioactive substance (0.5 micrograms of glycine 14C or methionine 35S) and colouring hair on the head. In a day or two labeled sousliks were caught again and in 6-7 days a total catching of animals and collection of ectoparasites from their hair and burrows done. In al, 942 sousliks were caught in the experimental areas and 2624 specimens of fleas of different species and 2305 ticks of R. schulzei collected. The average number of fleas which were labeled on one little souslik for a day was 0.1 to 2.3. The number of labeled fleas in the above areas coincided with the number of fleas infected with plague in the areas of epizootia. The level of parasitic contact in settlements of little souslik cannot provide the development of plague epizootia by transmission of the agent through fleas.     

Long-term egg retention and parasitization in Trichogramma principum (Hym., Trichogrammatidae)

Laboratory experiments with Trichogramma principium Sug. et Sor. females that were offered Sitotroga cerealella Oliv. eggs demonstrated that less than half of the ovipositing females started oviposition during the first 2 days of the experiment, whereas the rest of the ovipositing females showed a delay in parasitization ranging from 2 to 10 days after contact with the host. Almost 10% of the wasps refused to parasitize the grain moth eggs over 12 days. The delay in parasitization may be as long as 6–8 days without any significant decrease in the number of mature ovarial eggs, in the number of eggs laid during the first 48 h of oviposition, and in the total lifetime fecundity. This egg retention is responsible for the fact that in spite of a relatively short mean duration of the oviposition period in each individual female (approximately 4 days), host parasitization by a group of simultaneously emerged wasps was almost uniformly distributed over 8–10 days. When induced, the parasitization state (i.e. the tendency to parasitize sequentially offered portions of host eggs) was stable both in the presence of a host and under host deprivation extended up to 8 days. These data provide further evidence for our hypotheses that the stability of the parasitization state in Trichogramma is based on endocrine mechanisms.     

寄主植物对椰心叶甲生长发育和繁殖力的影响

研究了5种棕榈科寄主植物椰子(Cocos nucifera)、酒瓶椰子(Hyophorbe lageni-caulis)、老人葵(Washingtonia filifera)、大王棕(Roystonea regia)和槟榔(Areca catechu)未展开心叶对椰心叶甲实验种群生长发育和繁殖力的影响.结果表明:在取食不同的寄主植物心叶的情况下,椰心叶甲完成1个世代所需的时间差异显著,其中取食老人葵所需时间最长,为72.8d,取食酒瓶椰子所需时间最短,为39.8d;酒瓶椰子上的平均蛹质量较其它4种寄主植物大;取食老人葵的椰心叶甲平均产卵量最高,为157.6粒,而取食槟榔的最少,为65.2粒;取食酒瓶椰子的椰心叶甲成虫寿命为207.5d,远比寄生于其它寄主的成虫寿命长.5种寄主植物上的实验种群趋势指数分别为53.57、54.98、48.56、20.46和11.54.     

Behavioural responses to ectoparasites: time-budget adjustments and what matters to Blue Tits Parus caeruleus infested by fleas

Blue Tit nests are often heavily infested by fleas, which feed on the incubating female and the nestlings. Depending on habitat quality, the drawing of blood by fleas reduces offspring quality, or it is compensated by an increase in food provisioning by the adults and may reduce their future reproduction. Given these fitness costs, tits are expected to have evolved behavioural responses enabling them to remove, destroy or minimize the contact with fleas. To identify these traits, we video-recorded the changes in frequency and duration of the hosts' potential anti-flea behavioural defences in nests experimentally infested with low and high flea densities. We also investigated whether flea load affected the number of male feeds delivered to incubating females, and whether the parents increased their rate of food provisioning to the nestlings equally at high flea density. Flea density significantly affected the nest sanitation and sleeping behaviour of Blue Tit females but had no significant effect on grooming. Female Blue Tits increased the frequency but decreased the duration of bouts of these behavioural traits, and hence their time-budgets, based on per hour duration of behaviour, were not significantly affected by flea density. High flea density reduced nestling weight at the early nestling stage but these costs were fully compensated by an increase in female feeding effort. Males did not increase their frequency of food provisioning to incubating females nor to nestlings in heavily infested nests. The results are discussed in the light of parasite-mediated selection on host behaviour and the reciprocal host selection on flea life-history and behavioural traits.     

Sympatric and allopatric combinations of hen fleas and great tits: a test of the local adaptation hypothesis

To test whether allopatric nest parasites differ from sympatric ones in their effect on various life history traits of their avian host, I designed a cross-transfer experiment in which hen fleas (Ceratophyllus gallinae) were exchanged between great tit (Parus major) nests in two geographically widely separated areas. In neither of the areas did allopatric fleas influence body mass, tarsus length, wing length, duration of nestling period or mortality in great tit nestlings more severely than did sympatric fleas. Duration of incubation was also similar among females independent of experimental treatment. This lack of difference between allopatric and sympatric fleas is hypothesized to reflect the comparative harmlessness of hen fleas for their hosts.     

On the equivalence of host local adaptation and parasite maladaptation: an experimental test

In spatiotemporally varying environments, host-parasite coevolution may lead to either host or parasite local adaptation. Using reciprocal infestations over 11 pairs of plots, we tested local adaptation in the hen flea and its main host, the great tit. Flea reproductive success (number of adults at host fledging) was lower on host individuals from the same plot compared with foreign hosts (from another plot), revealing flea local maladaptation. Host reproductive success (number of fledged young) for nests infested by foreign fleas was lower compared with the reproductive success of controls, with an intermediate success for nests infested by local fleas. This suggests host local adaptation although the absence of local adaptation could not be excluded. However, fledglings were heavier and larger when reared with foreign fleas than when reared with local fleas, which could also indicate host local maladaptation if the fitness gain in offspring size offsets the potential cost in offspring number. Our results therefore challenge the traditional view that parasite local maladaptation is equivalent to host local adaptation. The differences in fledgling morphology between nests infested with local fleas and those with foreign fleas suggest that flea origin affects host resource allocation strategy between nestling growth and defense against parasites. Therefore, determining the mechanisms that underlie these local adaptation patterns requires the identification of the relevant fitness measures and life-history trade-offs in both species.