Experimental study of the infecting ability of the flea Coptopsylla lamellifer rostrata in the Kyzylkum natural focus of plague

Experimental infecting of the fleas Coptopsylla lamellifer rostrata by plague was carried out. The fleas were infected by feeding through biomembrane (skin of a white mouse) with a mixture of defibrinated blood of guinea-pig and plague microbes. Under concentration 2-3 milliard of the microbes in 1 millilitre, from 60 to 100% of the fleas were infected. Forming of the block of proventriculus was observed in 9-13 day. Mean percent of blocking for all experiments was less than 1%. Maximal rate of blocking (2.8%) was observed in the experiment under everyday 3-hour feeding. After planting of the fleas with blocks on great gerbil (2 fleas on one gerbil) 20% of the gerbils were infected. The conclusion had been made that C. lamellifer rostrata has only insignificant effectiveness as a vector of plague agent in Kyzylkum. Probably it is caused by some specific features of this subspecies.     

Evidence for a negative fitness-density relationship between parent density and offspring quality for two Xenopsylla spp. parasitic on desert mammals

We tested the hypothesis of negative fitness-density relationships and predicted that an increase in the density of parent fleas would result in lower survival rates and longer development time of pre-imagoes as well as shorter time to death from starvation of newly emerged imagoes. These predictions were experimentally tested on Xenopsylla conformis Wagner and Xenopsylla ramesis Rothschild feeding on two rodent species, Meriones crassus Sundevall or Dipodillus dasyurus Wagner. Survival of larvae and pupae, but not eggs, was negatively affected by parent density. An increase in parent density led to a decrease in the number of imagoes of the next generation. Eggs of both species developed faster when the parents were at low densities on either host. The same was true for larval X. ramesis, but not larval X. conformis. The negative effects of parent density on the duration of pupal development were evident in X. conformis, parents of which fed on both hosts, and X. ramesis from parents fed on M. crassus, whereas X. ramesis from parents fed on D. dasyurus developed faster at low densities. A negative effect of density on the development of offspring from egg to imago in X. conformis was manifested mainly when parent fleas fed on D. dasyurus, whereas the negative effect of density on offspring development in X. ramesis was manifested mainly when parent fleas fed on M. crassus. Although there was no general effect of parent density on the resistance of imago offspring to starvation, male X. ramesis from parents fed at the highest density survived starvation for significantly shorter times compared with those from parents fed at lower densities. Manifestation of the negative effect of parent density on offspring quality appears to vary with flea species and may be affected by host species.     

Fleas of the eastern Kyzyl Kum and their epizootic significance]

The fauna of fleas in the East Kisil-Kum includes 24 species. Different physicogeographic regions of sands are characterized by different spectra and number of species of fleas. The greatest share in collections is composed by Xenopsylla gerbilli caspica--the parasite of Rhombomys opimus. In autumn the other parasite of this rodent, Coptopsylla lamellifer, is a mass species. All the most significant and long changes in the number of these species are associated with changes in the density of their hosts' population. The most significant in epizootic respect is the flea X. gerbilli caspica from which about 50% of plague agent cultures were isolated.     

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.     

Larval interspecific competition in two flea species parasitic on the same rodent host

Abstract.  1. The fleas Xenopsylla conformis and Xenopsylla ramesis exploit the same rodent host, Meriones crassus , and replace each other between two different habitats situated at the opposite sides of a steep precipitation gradient. It was hypothesised that the reason for this paratopic distribution is competition between larvae of the two species for food resources.
2. This hypothesis was tested by studying the performance of larvae of the two species in terms of their developmental success in mixed-species and single-species treatments under different air temperatures, relative humidities, substrate textures, and food abundance.
3. The number of individuals of X. conformis that survived until emergence depended significantly on the presence of competing species, being, in general, lower in mixed-species compared with single-species treatments. The decrease in developmental success of X. conformis in mixed-species treatments was found mainly during food shortage. In contrast, presence of the competitor did not affect the number of X. ramesis that survived until emergence. No effect of the presence of the competitor on duration of development or sex ratio was found in either species.
4. The results of this study, together with the results of our previous studies, provide an explanation for the paratopic distribution of X. conformis and X. ramesis that exploit the same host species.     

A case of experimental transmission of the causative agent of plague from Ornithodoros tartakovskyi ticks to fleas of the great gerbil

Biological interrelations between ectoparasites from the colonies of great gerbil were studied experimentally. Hungry fleas Coptopsylla lamellifer are able to feed on the ticks Ornithodoros tartakovskyi infected with plague and in so doing be infected with plague agent. Such feeding is identified as "heterovampirism".     

Characteristics of digestion of blood of different animals by the flea Xenopsylla cheopis]

Under experimental conditions fleas X. cheopis engorged and assimilated blood of 4 species of rodents, man, pigeon and two species of reptiles. Histological investigations have shown that the structure of food clot, destruction rate of blood cells and general duration of digestion are changed considerably depending on the host's blood. In females the digestion of white mice blood to haematin at 23-24 degrees lasted not over 12 hours, the digestion of blood of hamsters and man - 12 to 20 hours, of guinea pigs and reptiles - 18 to 25 hours and of white rats and pigeons - from 20 to 30 hours. In males the digestion carries out somewhat slower. The above differences in blood digestion can apparently create unequal conditions for pathogenic microbes in the intestine of fleas that affects the preservation and reproduction of the agent in the organisms of these carriers.     

Density-dependent host selection in ectoparasites: An application of isodar theory to fleas parasitizing rodents

Parasites should make the same decisions that every animal makes regarding fitness reward. They can maximize reproductive success by selection of those habitats that guarantee the greatest fitness output. We consider the host population as a habitat of a parasite population. Consequently, hosts (=habitats) that differ quantitatively or qualitatively will support different numbers of parasites. The nature of habitat selection can be detected by isodars, lines along which habitat selection yields equivalent fitness reward. We applied this approach to study host selection of five fleas, each infesting two desert rodents. Xenopsylla conformis, Xenopsylla ramesis, Nosopsyllus iranus theodori and Stenoponia tripectinata medialis parasitize Gerbillus dasyurus and Meriones crassus. Synosternus cleopatrae pyramidis parasitizes Gerbillus andersoni allenbyi and Gerbillus pyramidum. Three fleas ( X. conformis, X. ramesis and S. c. pyramidis) were able to perceive quantitative (amount of the resource; e.g. organic matter in the nest for flea larvae) and/or qualitative (pattern of resource acquisition; e.g. host defensiveness) differences between hosts. Two other fleas did not perceive between-host differences. X. conformis was a density-dependent host selector that showed sharp selectivity at low density. X. ramesis and S. c. pyramidis were density-independent host selectors with a direct correspondence of density with habitat quality. N. i. theodori and S. t. medialis were non-selectors with no relationship at all between density and host quality. The results of the application of the isodar theory suggest that ectoparasites, like other animals, behave as if they are able to make choices and decisions that favour environments in which their reproductive benefit is maximized.     

Herpesvirus sylvilagus in cottontail rabbits: attempted laboratory transmission by two insect species.

The vector potential of the rabbit flea (Cediopsylla simplex) and a mosquito (Aedes triseriatus) was investigated for Herpesvirus sylvilagus transmission among cottontail rabbits (Sylvilagus floridanus). Twelve groups of 12-50 fleas were fed on three viremic cottontails for 2-21 days before transfer to 12 susceptible rabbits. Standard interrupted feeding trials employed five groups of 6-12 mosquitoes, two viremic donor cottontails anf five healthy recipients. No evidence of virus was detected from recipients' blood nor did they develop specific antibody. Virus acquisition and persistence in the insects was evaluated by attempting to recover the virus from 19 pools of mosquitoes engorged on viremic blood and 36 pools of engorged fleas or those living on viremic hosts for 1-21 days. Results were negative.     

Characterization of the salivary apyrase activity of three rodent flea species

1. Salivary gland lysates of the adult female fleas Oropsylla bacchi, Orchopea howardi and Xenopsylla cheopis hydrolyse ATP and ADP, but not AMP, thus characterizing the existence of a salivary apyrase activity. 2. In all species Mg++ or Ca++ function as activators, and a pH optimum between 7 and 8 is observed. 3. Salivary gland lysates of male fleas contain significantly smaller amounts of the enzyme activity than do those of female fleas. 4. Immediately following a blood meal, apyrase activity and protein content of female X. cheopis salivary glands are 2-3-fold less than that of unfed fleas, indicating that salivary apyrase activity is secreted during feeding. 5. It is suggested that, as in other arthropods, salivary apyrase may facilitate blood location and blood feeding by preventing ADP-induced platelet aggregation at the site of the bite.     

Metabolic rate and jump performance in seven species of desert fleas

We hypothesized that sexual and interspecific differences in jumping performance of fleas found in our previous study are correlated with differences in resting metabolic rate (RMR) between sexes and among species. To test this hypothesis, we measured RMR of seven flea species (Xenopsylla conformis mycerini, Xenopsylla ramesis, Xenopsylla dipodilli, Parapulex chephrenis, Synosternus cleopatrae pyramidis, Nosopsyllus iranus theodori and Stenoponia tripectinata medialis). We compared RMR between sexes and among species and examined whether there is intra- and interspecific correlation between RMR and jumping ability. Both mass-specific and mass-independent RMR were the highest in female S. t. medialis, whereas mass-specific RMR was the lowest in male X. dipodilli and mass-independent RMR was the lowest in three Xenopsylla species and P. chephrenis. Mass-specific and mass-independent RMR were significantly higher in females than in males in all fleas except S. t. medialis. Differences in jumping ability between males and females were found to be correlated with sexual differences in mass-specific or mass-independent RMR. Interspecific comparison showed that the length of jump in both male and female fleas was strongly affected by their mass-specific and mass-independent RMR.     

长爪沙鼠体蚤和巢蚤数量研究

1982～1996年对内蒙古自治区鄂托克前旗长爪沙鼠Meriones unguiculatus体蚤和巢蚤的数量进行了调查和分析,得到如下结果。获体蚤15种,同型客蚤指名亚种Xenopsylla conformis conformls (50.8%) 和秃病蚤蒙冀亚种Nosopsyllus laveiceps kuzenkovi (40.6%)为优势种。获巢蚤15种,秃病蚤蒙冀亚种(74.3%)为优势种;盔状新蚤Neopsylla galea (11.9%)和叶状切唇蚤突高亚种Coptopsyllus lamelliferardua (8.1%)为常见种。年巢蚤指数的均值是年体蚤指数的6.92倍。体蚤指数与巢蚤指数不相关(P>0,05),体染蚤率与巢染蚤率不相关(P>0.05)。不同年份 的体蚤和巢蚤多样性比较稳定。连续两年春季或秋季秃病蚤巢蚤指数大于10只后,可能流行动物鼠疫。     

Polymerase chain reaction (PCR) identification of rodent blood meals confirms host sharing by flea vectors of plague

Elucidating feeding relationships between hosts and parasites remains a significant challenge in studies of the ecology of infectious diseases, especially those involving small or cryptic vectors. Black‐tailed prairie dogs (Cynomys ludovicianus) are a species of conservation importance in the North American Great Plains whose populations are extirpated by plague, a flea‐vectored, bacterial disease. Using polymerase chain reaction (PCR) assays, we determined that fleas (Oropsylla hirsuta) associated with prairie dogs feed upon northern grasshopper mice (Onychomys leucogaster), a rodent that has been implicated in the transmission and maintenance of plague in prairie‐dog colonies. Our results definitively show that grasshopper mice not only share fleas with prairie dogs during plague epizootics, but also provide them with blood meals, offering a mechanism by which the pathogen, Yersinia pestis, may be transmitted between host species and maintained between epizootics. The lack of identifiable host DNA in a significant fraction of engorged Oropsylla hirsuta collected from animals (47%) and prairie‐dog burrows (100%) suggests a rapid rate of digestion and feeding that may facilitate disease transmission during epizootics but also complicate efforts to detect feeding on alternative hosts. Combined with other analytical approaches, e.g., stable isotope analysis, molecular genetic techniques can provide novel insights into host‐parasite feeding relationships and improve our understanding of the role of alternative hosts in the transmission and maintenance of disease.     

Facultative imaginal diapause in Xenopsylla conformis (Siphonaptera) fleas

The factors favouring the cessation of reproduction in X. conformis are laid in the preimaginal state. The drop in temperature during the formation of imago at the pupal stage is a signal for the cessation of reproduction. Imagos hatched at a temperature lower than that of developmental conditions of preimaginal stages do not start reproduction and enter facultative imaginal diapause state. With further decrease in temperature the state of fleas intensifies. With the rise of temperature fleas come out of diapause. In autumn coming out of diapause begins at a temperature higher than 20 degrees, on the 8th--9th day. The lower air temperature the more rapid is coming out of diapause, at a rise of temperature of 3 to 5 degrees.     

Ectoparasite fitness in auxiliary hosts: phylogenetic distance from a principal host matters

We studied reproductive performance in two flea species (Parapulex chephrenis and Xenopsylla ramesis) exploiting either a principal or one of eight auxiliary host species. We predicted that fleas would produce more eggs and adult offspring when exploiting (i) a principal host than an auxiliary host and (ii) an auxiliary host phylogenetically close to a principal host than an auxiliary host phylogenetically distant from a principal host. In both flea species, egg production per female after one feeding and production of new imago after a timed period of an uninterrupted stay on a host differed significantly between host species. In general, egg and/or new imago production in fleas feeding on an auxiliary host was lower than in fleas feeding on the principal host, except for the auxiliary host that was the closest relative of the principal host. When all auxiliary host species were considered, we did not find any significant relationship between either egg or new imago production in fleas exploiting an auxiliary host and phylogenetic distance between this host and the principal host. However, when the analyses were restricted to auxiliary hosts belonging to the same family as the principal host (Muridae), new imago production (for P. chephrenis) or both egg and new imago production (for X. ramesis) in an auxiliary host decreased significantly with an increase in phylogenetic distance between the auxiliary and principal host. Our results demonstrated that a parasite achieves higher fitness in auxiliary hosts that are either the most closely related to or the most distant from its principal host. This may affect host associations of a parasite invading new areas.     

Sex ratio in flea infrapopulations: number of fleas, host gender and host age do not have an effect

This study set out to determine whether the sex ratio of fleas collected from host bodies is a reliable indicator of sex ratio in the entire flea population. To answer this question, previously published data on 18 flea species was used and it was tested to see whether a correlation exists between the sex ratio of fleas collected from host bodies and the sex ratio of fleas collected from host burrows. Across species, the female:male ratio of fleas on hosts correlated strongly with the female:male ratio of fleas in their burrows, with the slope of the regression overlapping 1. Controlling for flea phylogeny by independent contrasts produced similar results. It was also ascertained whether a host individual is a proportional random sampler of male and female fleas and whether the sex ratio in flea infrapopulations depends on the size of infrapopulations and on the gender and age of a host. Using field data, the sex ratio in infrapopulations of 7 flea species parasitic on 4 rodent species was analysed. Populations of 3 species (Nosopsyllus iranus, Parapulex chephrenis and Xenopsylla conformis) were significantly female-biased, whereas male bias was found in 1 species (Synosternus cleopatrae). In general, the sex ratio of fleas collected from an individual rodent did not differ significantly from the sex ratio in the entire flea population. Neither host gender, and age nor number of fleas co-occurring on a host affected (a) the sex ratio in flea infrapopulations and (b) the probability of an infrapopulation to be either female- or male-biased.     

Productivity of cultures of 4 species of fleas in laboratory breeding

When rearing fleas under laboratory conditions maximum productivity was observed in breeding cages with 100 fleas (80 females and 20 males). When the number of fleas was increased to 250 and 500 specimens the progeny decreased in number. At the ratio between males and females of 1 : 2 and 1 : 4 the productivity of fleas virtually did not change. Of all species studied fleas of X. cheopis are characterized by most high productivity. In all months of the year the average number of hatched fleas per one cage was 13-15 thou. of X. cheopis, 7-8 thou. of C. tesquorum, 1.5-2.5 thou. of X. skrjabini, and 3-4 thou. of Ct. wladimiri.     

中国云南部分人间鼠疫流行区蚤类区系调查(英文)

归纳了中国云南 13个人间鼠疫流行区的调查资料 ,对调查疫区的蚤类区系进行了研究。总计捕获12 0 77只小兽 ,隶属啮齿目、食虫目及攀目 3个目中的 9科、2 9属、4 7种。从小兽体表共采获 9369只蚤 ,经分类鉴定 ,隶属 5科、18属、33种。 33种蚤及 4 7种小兽宿主均按其分类阶元详细列于文末。结果表明 ,山区蚤及小兽宿主的种数明显多于坝区。坝区农耕地的优势种相对简单 ,优势种地位突出 ,黄胸鼠及印鼠客蚤分别是最重要的宿主及蚤种 (构成比分别为 83 2 7%和 75 32 % )。山区的优势种相对较复杂 ,优势种的种类较多 ,但其构成比较低 ( 10 96%～ 4 7 95% )。黄胸鼠及绒鼠为山区地带的两种优势宿主 ,缓慢细蚤、端凹栉眼蚤、印鼠客蚤、偏远古蚤及短突栉眼蚤为山区地带的 5种优势蚤种。多数蚤种可寄生两种以上的小兽宿主 ,但其所寄生的主要宿主并不多。结果提示 ,作为疫区主要媒介的印鼠客蚤及其所对应的主要寄生宿主 (黄胸鼠 )在坝区突出的优势种地位 ,似可解释近年疫区的鼠疫病人主要出现在坝区的原因     

Respiratory gas exchange in the desert flea Xenopsylla ramesis (Siphonaptera: Pulicidae): response to temperature and blood-feeding

Xenopsylla ramesis is a flea species parasitizing gerbilline rodents in the deserts of the Middle East. This study was undertaken to determine metabolic requirements of the different developmental stages of the flea-life cycle as well as to investigate the metabolic response to temperature and starvation after blood feeding. A high resolution respirometry system was used to measure CO2 emission of fleas ranging in size from 0.166+/-0.006 mg (larvae) to 0.263+/-0.009 mg (adults). The free-living stages (larvae and adults) had significantly higher metabolic rates than the cocooned stages (pupae). CO2 emission rates of the larvae exceeded that of the adults by 2.6-fold and the pupae by 7.3 times. In the adults, both temperature and blood feeding significantly affected starvation-level metabolism. Metabolism was temperature dependent with an average Q10 of 2.57 for females and 2.55 for males over the temperature range of 10-30 degrees C. No consistent decline in thermal sensitivity at higher ambient temperatures was evident. Fleas that had a blood meal prior to starvation had significantly higher metabolic rates (0. 86 +/- 0.008 x 10(-3) ml mg(-1) h(-1)) than fleas, which were newly emerged unfed adults (0.56 +/- 0.1 x 10(-3) ml mg(-1) h(-1)). Water content also differed between fed (range approx. 67-69% body mass) and newly emerged adults (range approx. 73-75% of body mass). Feeding may stimulate some as yet undetermined physiological process that causes differential metabolic response in starving, fed and unfed fleas. Characteristics of gas exchange in desert-dwelling fleas are reflective of the off-host life style in the protected microenvironment of the host nest or burrow, rather than as a response to any type of environmental extreme.