Wolbachia endosymbionts in fleas (Siphonaptera)

Intracellular endosymbionts, Wolbachia spp., have been reported in many different orders of insects and in nematodes but not previously in fleas. This is the first conclusive report of Wolbachia spp. within members of the Siphonaptera. Using nested polymerase chain reaction (PCR) targeting of the 16S ribosomal RNA gene, we screened for Wolbachia spp. in fleas collected from 3 counties in Georgia and 1 in New York. The prevalence of Wolbachia spp. detected varied among the 6 different species screened: 21% in the cat flea Ctenocephalides felis (n = 604), 7% in the dog flea C. canis (n = 28), 25% in Polygenus gwyni (n = 8), 80% in Orchopeas howardi (n = 15), 94% in Pulex simulans (n = 255), and 24% in the sticktight flea Echidnophaga gallinacea (n = 101). Wolbachia spp. infection in fleas was confirmed by sequencing positive PCR products, comparing sequenced 16S ribosomal DNA (rDNA) with Wolbachia spp. sequences in GenBank using BLAST search, and subjecting sequence data to phylogenetic analysis. For further confirmation, 16S rDNA-positive samples were reamplified using the wsp gene.     

The potential role of swift foxes (Vulpes velox) and their fleas in plague outbreaks in prairie dogs

Swift foxes (Vulpes velox) have been proposed as potential carriers of fleas infected with the bacterium Yersinia pestis between areas of epizootics in black-tailed prairie dogs (Cynomys ludovicianus). We examined antibody prevalence rates of a population of swift foxes in Colorado, USA, and used polymerase chain reaction (PCR) assays to examine their flea biota for evidence of Y. pestis. Fifteen of 61 (24%) captured foxes were seropositive, and antibody prevalence was spatially correlated with epizootic plague activity in prairie dog colonies in the year of, and previous to, the study. Foxes commonly harbored the flea Pulex simulans, though none of the fleas was positive for Y. pestis.     

Host associations and origin in the formation of the Caucasian fauna of fleas (Siphonaptera)

Results of analysis of the Caucasian fauna of fleas and their association with mammal and avian hosts are reported. The Caucasian fauna of potential flea hosts comprises about 130 species of mammals and about 470 species of birds. Most of the flea species in the Caucasian fauna (88 out of 155) parasitize rodents, 51 species of which are permanent hosts of different flea species; 13 flea species occur on 11 species of insectivores; 13 flea species, on 13 species of chiropterans; 14 flea species, on 20 species of carnivores. Only 2 flea species parasitize artiodactyles. 54 species of birds are permanent hosts of 23 species of fleas from 4 genera in the Caucasus. Ten types of ranges of flea species are distinguished; host associations of the Caucasian flea species from these groups are discussed. The greatest numbers of hosts from the families Cricetidae, Muridae, and Sciuridae are associated with fleas with Euro-Asian (extra-Siberian), European, Turanian, and Iranian ranges. Soricidae are known as hosts of flea species with European and Euro-Turanian ranges. Four major groups of flea taxa are represented in the Caucasian fauna. The distribution of the first group is determined by the influence of the palaeofauna of the ancient European continent in the early Cenozoic; that of the second group, by the influence of the fauna of the ancient Asian continent during the Paleogene and part of the Neogene; the third, by the influence of the fauna of southern Europe starting with the Miocene. The fourth group comprises the species which immigrated from northern Europe and Asia in the Late Neogene (2–3 mln years ago).     

Epizootiology of Haemoproteus danilewskyi (Haemosporina: Haemoproteidae) in blue jays (Cyanocitta cristata) in southcentral Florida

Prevalence and density of Haemoproteus danilewskyi was studied in a population of free-ranging blue jays (Cyanocitta cristata) in southcentral Florida (USA) from May 1992 to December 1995. Prevalence of infection was 27% for data combined over years, seasons, ages, and sexes. Prevalence did not vary between sexes or among years, but increased with age and varied with season, being highest in June-July and lowest in November-January. Parasite density did not vary between sexes or among seasons, but was higher in younger birds when controlling for season. To determine periods of natural transmission, seasonal patterns of infection were compared with previous month abundance of the biting fly vectors. Mean monthly prevalence of H. danilewskyi in older jays was positively correlated with previous month abundance of Culicoides edeni and C. arboricola, both capable of sporogonic development of H. danilewskyi.     

Systemic efficacy of nodulisporamides against fleas on dogs.

Nodulisporic acid A (NSA) has been shown previously to be safe in dogs and to deliver >90% flea control for 4 days following a single oral administration. Three newly prepared nodulisporamide derivatives were subsequently identified from an artificial membrane flea feeding system as exhibiting potency substantially greater than NSA. To determine if they have superior in vivo activity, these 3 nodulisporamides, as well as NSA, were evaluated in dogs at 15 mg/kg/os. Parasite challenges were made by placing 100 live Ctenocephalides felis fleas onto the dorsum of dogs every 48 hr and examining efficacy at each of those intervals over a 22-day period. Results showed that NSA produced >90% efficacy at day 2 and 81% efficacy at day 4, and its residual flea killing fell to approximately 50% by day 6 posttreatment. All dogs treated with the 3 new experimental nodulisporamides were 100% protected from flea challenges to day 8 posttreatment, and 2 of the compounds continued to produce >90% residual activity to 2 wk posttreatment. Pharmacokinetic analysis showed that plasma profiles and half-lives of NSA and these 3 new compounds correlated closely with flea efficacy. These results demonstrate that specific substitutions to the pharmacophore of NSA can substantially increase the duration of activity against fleas.     

Sampling fleas: the reliability of host infestation data

The use of measures of host infestation as a reliable indicator of a flea population size to be used in interspecific comparisons was considered. The abundance of fleas collected from host bodies and collected from host burrows was compared among 55 flea species, controlling for the effect of flea phylogeny. The mean number of fleas on host bodies correlated positively with the mean number of fleas in host burrows/nests both when the entire data pool was analysed and for separate subsets of data on 'fur' fleas and 'nest' fleas. This was also true for a within-host (Microtus californicus) between-flea comparison. The results of this study demonstrate that, in general, the index of host body infestation by fleas can be used reliably as an indicator of the entire population size.     

Feeding performance of fleas on different host species: is phylogenetic distance between hosts important?

We asked if and how feeding performance of fleas on an auxiliary host is affected by the phylogenetic distance between this host and the principal host of a flea. We investigated the feeding of 2 flea species, Parapulex chephrenis and Xenopsylla ramesis, on a principal (Acomys cahirinus and Meriones crassus, respectively) and 8 auxiliary host species. We predicted that fleas would perform better (higher proportion of fleas would feed and take larger bloodmeals) on (a) a principal rather than an auxiliary host and (b) auxiliary hosts phylogenetically closer to a principal host. Although feeding performance of fleas differed among different hosts, we found that: (1) fleas did not always perform better on a principal host than on an auxiliary host; and (2) flea performance on an auxiliary host was not negatively correlated with phylogenetic distance of this host from the principal host. In some cases, fleas fed better on hosts that were phylogenetically distant from their principal host. We concluded that variation in flea feeding performance among host species results from interplay between (a) inherent species-specific host defence abilities, (b) inherent species-specific flea abilities to withstand host defences and (c) evolutionary tightness of association between a particular host species and a particular flea species.     

Aggregation and species coexistence in fleas parasitic on small mammals

The aggregation model of coexistence states that species coexistence is facilitated if interspecific aggregation is reduced relative to intraspecific aggregation. We investigated the relationship between intraspecific and interspecific aggregation in 17 component communities (the flea assemblage of a host population) of fleas parasitic on small mammals and hypothesized that interspecific interactions should be reduced relative to intraspecific interactions, facilitating species coexistence. We predicted that the reduction of the level of interspecific aggregation in relation to the level of intraspecific aggregation would be positively correlated with total flea abundance and species richness of flea assemblages. We also expected that the higher degree of facilitation of flea coexistence would be affected by host parameters such as body mass, basal metabolic rate (BMR) and depth and complexity of burrows. Results of this study supported the aggregation model of coexistence and demonstrated that, in general, a) conspecific fleas were aggregated across their hosts; b) flea assemblages were not dominated by negative interspecific interactions; and c) the level of interspecific aggregation in flea assemblages was reduced in relation to the level of intraspecific aggregation. Intraspecific aggregation tended to be correlated positively to body mass, burrow complexity and mass-independent BMR of a host. Positive interspecific associations of fleas tended to occur more frequently in species-rich flea assemblages and/or in larger hosts possessing deep complex burrows. Intraspecific aggregation increased relative to interspecific aggregation when species richness of flea infracommunities (the flea assemblage of a host individual) and component communities increased. We conclude that the pattern of flea coexistence is related both to the structure of flea communities and affinities of host species.     

云南省玉龙鼠疫疫源地野外鼠形动物寄生蚤丰盛度影响因素分析

【目的】分析云南省玉龙鼠疫疫源地野外鼠形动物寄生蚤丰盛度的影响因素。【方法】选取云南省玉龙鼠疫疫源地3个海拔区域,按4个季节进行野外捕鼠,捕获的鼠形动物用梳检法收集体表寄生蚤并在显微镜下分类鉴定。通过实际测量和实地观察相结合的方式收集潜在影响鼠形动物寄生蚤丰盛度的因素包括鼠形动物特征变量指标(如种类、年龄、性别、体长、体重)、环境和气象因子(如海拔、季节)等数据。采用EpiData 3.02软件建立数据库,在R软件下使用跨栏负二项分布回归分析鼠形动物寄生蚤丰盛度的影响因素。【结果】从捕获的884只鼠形动物中检获寄生蚤9种484头,以特新蚤指名亚种、方叶栉眼蚤、无值大锥蚤、云南栉眼蚤为主(86.16%)。回归分析显示： 2 700-3 000 m和3 000 m以上海拔鼠形动物染蚤概率较2 400-2 700 m分别增加1.27和3.72倍;湿度高于70%时,鼠形动物染蚤概率较湿度≤70%时减少41%;与齐氏姬鼠的染蚤概率相比,中华姬鼠的染蚤概率降低50%,大绒鼠的染蚤概率增加79%;体长超过104 mm的鼠形动物染蚤概率较体长≤104 mm的鼠形动物染蚤概率增加76%;气温高于15℃时,鼠形动物染蚤数量较温度≤15℃时降低67%;成年鼠形动物的染蚤数量较未成年鼠形动物的染蚤数量增加2.25倍;与春季相比,夏季的染蚤数量增加2.00倍,秋季的染蚤概率减少48%,冬季的染蚤概率和染蚤数量分别增加1.44和1.06倍。【结论】玉龙鼠疫疫源地野外鼠形动物寄生蚤以特新蚤指名亚种、方叶栉眼蚤、无值大锥蚤、云南栉眼蚤为优势蚤种。鼠形动物寄生蚤丰盛度与海拔、季节、气温、湿度等环境气象因子及鼠形动物种类、体长、年龄等鼠形动物特征变量密切相关。     

Mammal and flea relationships in the Great Basin Desert: from H. J. Egoscue's collections

Host-parasite association among 58 flea species parasitizing 40 mammal species in the Great Basin Desert of the western United States was investigated. Increased flea species richness was correlated with larger geographic ranges and stable locomotion of hosts. Hosts from habitats of moderately low productivity (sage and grass) and of Peromyscus maniculatus size, 10-33 g, had the highest flea species richness. Larger hosts had fewer flea species, but fleas were more prevalent. Increased host species richness correlated with flea species eye size. Mammals clustered into 3 major and 1 minor ecological groups, and fleas clustered into 2 major groups among rodents, and 6 minor groups, forming 12 host-parasite biocenoses. Factors producing biocenoses were shared burrows of mice and rats; food chains of hares, rabbits, squirrels, and their predators; keystone mammals: Lagurus curtatus, Neotoma lepida, Ochotona princeps, and Spermophilus townsendii; keystone fleas: Megabothris abantis, and Meringis hubbardi; or host isolation, Neotoma cinerea with Oropsylla montana, Sorex vagrans with Corrodopsylla curvata, and Tadarida brasiliensis with Sternopsylla distincta. Although host relatedness accounted for flea prevalence, host sociality explained the presence or absence of mammal-flea relationships.     

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.     

Host distribution and pathogen infection of fleas (Siphonaptera) recovered from small mammals in Pennsylvania

The number of recognized flea‐borne pathogens has increased over the past decade. However, the true number of infections related to all flea‐borne pathogens remains unknown. To better understand the enzootic cycle of flea‐borne pathogens, fleas were sampled from small mammals trapped in central Pennsylvania. A total of 541 small mammals were trapped, with white‐footed mice (Peromyscus leucopus) and southern red‐backed voles (Myodes gapperi) accounting for over 94% of the captures. Only P. leucopus were positive for examined blood‐borne pathogens, with 47 (18.1%) and ten (4.8%) positive for Anaplasma phagocytophilum and Babesia microti, respectively. In addition, 61 fleas were collected from small mammals and tested for pathogens. Orchopeas leucopus was the most common flea and Bartonella vinsonii subspecies arupensis, B. microti, and a Rickettsia felis‐like bacterium were detected in various flea samples. To the best of our knowledge, this is the first report of B. microti DNA detected from a flea and the first report of a R. felis‐like bacterium from rodent fleas in eastern North America. This study provides evidence of emerging pathogens found in fleas, but further investigation is required to resolve the ecology of flea‐borne disease transmission cycles.     

Distribution of fleas (Siphonaptera) among small mammals: mean abundance predicts prevalence via simple epidemiological model

We used data on the abundance and distribution of fleas parasitic on small mammals in Slovakia and aimed: (i) to confirm a positive relationship between abundance and distribution fleas within and across host species; and (ii) to test if prevalence of fleas can be reliably predicted from a simple epidemiological model that takes into account flea mean abundance and its variance. Prevalence of a flea species increased with an increase in its mean abundance both within and across host species. We calculated prevalences both for each flea-host association and for each flea species across all hosts. Observed prevalences did not differ significantly from those predicted by the epidemiological model using parameters of Taylor's power relationship between mean abundance of fleas and its variance. Regressions of predicted prevalences against observed prevalences produced slope values that did not differ significantly from unity and were independent of scale (within or across host species). Our results demonstrated that up to 96% of variance in flea prevalence can be explained solely by their mean abundance. We concluded that, in general, there is no need to invoke other, more complex factors for the explanation of the variation in flea prevalence.     

Geographic variation in rodent-flea relationships in the presence of black-tailed prairie dog colonies

We characterized the relationship between fleas and their rodent hosts in the presence of prairie dog colonies and compared them to adjacent assemblages away from colonies. We evaluated the rodent-flea relationship by quantifying prevalence, probability of infestation, flea load, and intensity of fleas on rodents. As prairie dog burrows provide refugia for fleas, we hypothesized that prevalence, flea load, and intensity would be higher for rodents that are associated with black-tailed prairie dog colonies. Rodents were trapped at off- and on-colony grids, resulting in the collection of 4,509 fleas from 1,430 rodents in six study areas. The rodent community composition varied between these study areas. Flea species richness was not different between prairie dog colonies and the surrounding grasslands (p = 0.883) but was positively correlated with rodent species richness (p = 0.055). Prairie dog colonies did not increase the prevalence of fleas (p > 0.10). Flea loads on rodents did not vary between off- and on-colony grids at three of the study areas (p > 0.10). Based on the prevalence, infestation rates, and flea loads, we identified Peromyscus maniculatus, Onychomys leucogaster, and two Neotoma species as important rodent hosts for fleas and Aetheca wagneri, Orchopeus leucopus, Peromyscopsylla hesperomys, Pleochaetis exilis, and Thrassisfotus as the most important fleas associated with these rodents. Prairie dog colonies did not seem to facilitate transmission of fleas between rodent hosts, and the few rodent-flea associations exhibited significant differences between off- and on-colony grids.     

The Effect of Initial Infection of <Emphasis Type="Italic">Citellophilus tesquorum altaicus</Emphasis> Ioff, 1936 (Siphonaptera: Ceratophyllidae) with <Emphasis Type="Italic">Yersinia pestis</Emphasis> on Its Alimentary Activity,Mortality, and Biofilm Formation

Alimentary activity and mortality was assessed in fleas Citellophilus tesquorum altaicus non-infected with Yersinia pestis and those with initial infection levels 50 and 100% during feeding on a non-specific host (white mice). The presence of the plague pathogen in fleas significantly stimulated their feeding activity, especially in females. No effect of infection on flea mortality was observed. At the same time, male fleas died more frequently than females.     

Australia’s vanishing fleas (Insecta: Siphonaptera): a case study in methods for the assessment and conservation of threatened flea species

While invertebrate conservation is attracting increased funding and interest, research remains heavily skewed towards ‘flagship’ insect groups like bees and butterflies. This has resulted in a knowledge gap relating to less popular but equally imperilled groups like fleas. Methods for the risk assessment of host specific parasites were used to determine the conservation status of all host specific flea species distributed in Australia. The results indicated one species apparently extinct, two critically endangered, two endangered, and three vulnerable. Based on these results, novel methods for the conservation of threatened fleas are outlined, including the concepts of holistic conservation and the cryptic loss effect.     

Geographical range size and host specificity in ectoparasites: a case study with Amphipsylla fleas and rodent hosts

Aim  To identify the factors that determine the geographical range sizes of ectoparasites with different degrees of host specificity.
Location  The study used data on the distributions of fleas of the genus Amphipsylla and their rodent hosts across the Holarctic.
Methods  All known points of occurrence of 32 flea species and 51 species of their rodent hosts were mapped. The shape and size of the geographical range of each species were estimated using a combination of the minimal convex polygon technique and modelling with the garp algorithm. Factors determining the geographical range sizes of the fleas were identified using stepwise multiple regression analysis.
Results  The geographical range size of fleas that are strongly host-specific across their entire ranges correlated positively with the geographical range size of the fleas' principal hosts, and negatively with the geographical range size of the fleas' potential competitors. The geographical range sizes of both (1) fleas that are locally host-specific but that shift their host preferences geographically, and (2) host-opportunistic fleas were positively correlated only with the area of the geographical ranges of their principal hosts. Strongly host-specific fleas occupied 0.2–80.0% of the geographical range of their principal hosts, whereas this figure was 0.9–83.7% in locally host-specific fleas and 16.6–63.7% in host-opportunistic fleas.
Main conclusions  The main determinant of the geographical range size of a flea species is the size of the geographical range of its hosts. The role of potential competitors in determining the geographical range size is stronger in host-specific than in host-opportunistic fleas. Cases in which the geographical range of a parasite is smaller than the geographical range(s) of its host(s) owing to narrower parasite environmental tolerances are much more frequent in host-opportunistic than in host-specific fleas.     

The flea epifauna of a suburban fox (Vulpes vulpes) population

Five hundred and thirteen fleas, of eight different species, were collected from a sample of 252 foxes killed in suburban London. 25–8% of foxes carried fleas, with a mean of 204 fleas per fox. Levels of infestation of male and female hosts did not differ significantly. Possible sources of the fleas infesting foxes are discussed with respect to their seasonal occurrence and fox prey composition. No evidence was found to support the suggestion that foxes obtain the majority of their fleas from prey items, although occasional heavy infestations of some flea species were probably derived from recent meals. Although Pulex irritans, Paraceras m. melis and Ctenocephalides canis , which contributed 35 % of the flea epifauna, could be considered parasitic on the fox, it seems probable that foxes pick up the majority of their fleas from the habitat through which they move. Thus, two particularly heavily infested categories of foxes were found: (1) juveniles during July-September, their fleas probably being accumulated during exploratory and play activities, and (2) all animals during the period October-December.     

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。     

Florida Scrub-Jays (Aphelocoma coerulescens) are Sentinels More When Well-Fed (Even with no Kin Nearby)

Sentinels occupy high, exposed positions while other group members forage nearby. If sentinel behavior involves a foraging–predation risk trade‐off, animals should be sentinels more when fed supplemental food. When individual Florida scrub‐jays (Aphelocoma coerulescens) were fed fragments of peanuts, during the following 30 min they shifted 30% of their time from foraging to sentinel behavior. In a follow‐up experiment, we fed either one or two members in each group. As before, the jays reduced their foraging and spent much more time as sentinels when given supplemental food. In each treatment, pairs were sentinels simultaneously considerably less often than expected by chance. The dramatic shift from foraging to sentinel behavior suggests that for Florida scrub‐jays sentinel behavior brings substantial benefits for no greater cost than that of lost opportunities to forage. Because the results held for simple mated pairs of scrub‐jays, we argue that kin selection and social prestige are not necessary to explain sentinel behavior.