Patterns of Formation of the Fauna of Fleas (Siphonaptera) Parasitizing Mammals in the Stavropol Upland and Adjacent Territories

According to the recent data, 55 species of fleas parasitizing 65 species of mammals were recorded in the Stavropol Upland (central Ciscaucasia, Russia). The fauna of the region comprises 87% of the total species composition of Ciscaucasian fleas. Most of these species are widely distributed on the Caucasian Isthmus and also in the Eastern Mediterranean. Of them, 14 species are polyzonal, 14 mostly occur in forest-steppes, 7 in foreststeppes and steppes, 12 in steppes and semi-deserts, and 7 species occur in semi-deserts. According to paleogeographic reconstructions, in the Pliocene the Stavropol Upland was colonized by the flea species from the forest landscapes of Southern Europe and also of Southwest Asia and Asia Minor. Later, in the Pleistocene and Holocene, the Caucasian Isthmus continued to be colonized by mesophilous species from Southern Europe and by semi-desert species from Southern Siberia and the Turanian Province.     

Fleas (Siphonaptera) associated with mammals and birds in the Ciscaucasia

The flea fauna of the Ciscaucasia comprises 76 species, 13 of which are associated with birds and the rest, with mammals. Rodent parasites are the most numerous; fleas associated with carnivores, bats, and insectivores are less abundant. Fleas parasitize different species of birds of the orders Passeriformes, Anseriformes, Falconiformes, and Strigiformes. Among 41 flea genera known from the Caucasus, species of the genera Amalaraeus, Araeopsylla, Atyphloceras, Caenopsylla, Callopsylla, Doratopsylla, Paraneopsylla, Peromyscopsylla, Phaenopsylla, Tarsopsylla, and Wagnerina are absent in the Ciscaucasia. Only two subendemic species were revealed in the region; 33 flea species are distributed over the entire Ciscaucasia, while the distribution of others is limited to landscapes of one or two natural zones.     

The Palaearctic centers of taxonomic diversity of fleas (Siphonaptera)

The Palaearctic flea fauna includes 921 species and 479 subspecies from 96 genera of 10 families. Of them, 858 species (94%) from 43 genera are endemic to the Palaearctic; they comprise 40% of the Palaearctic Hystrichopsyllidae, 24% of Ceratophyllidae, and 20% of Leptopsyllidae. Ranges of 581 species (63% of the Palaearctic fauna) are situated within one province or subregion of the Palaearctic. Species with ranges including a part of Asia (592) comprise 87% of the total fauna; 72% of the species (517) are endemic to the Palaearctic. The largest centers of taxonomic diversity of Palaearctic fleas are situated in the East Asian, Central Asian, and Turano-Iranian Subregions: 320 species of fleas (214 of them endemic) from 59 genera (8 endemic) are known from the East Asian Subregion; 270 species (over 120 endemic) from 54 genera (5 endemic) are distributed in the Central Asian Subregion. The Turano-Iranian fauna comprises 213 species (103 endemic) from 47 genera (3 endemic); about 160 species occur in the Turanian Subprovince closest to the Russian borders, one-third of them (52 species, or 33%) are endemic; 69 species more are endemic to the entire Asian part of the Palaearctic. Extra-Asian and extra-Siberian ranges are known in 190 flea species. In the western Palaearctic, 76 species are endemic to the European Province, and 57 species, to the Mediterranean Province; 36 species have Euro-Mediterranean distribution. The fauna of the Saharo-Arabian Subregion comprises 30 species (12 endemic), 6 species have ranges of the Mediterranean-Saharo-Arabian type. Scenarios of the origin of the Siphonaptera at the Triassic-Jurassic boundary are hypothesized. Formation of the Palaearctic flea fauna was mostly supported by the Asian-Indo-Malayan and East Asian-Western American palaeofaunal centers of taxonomic diversity. The long history of faunal exchange between the east Palaearctic and the west Nearctic is manifested by the distribution of the parasites of rodents and insectivores, fleas of the genera Stenoponia, Rhadinopsylla, Nearctopsylla, and Catallagia, belonging to several subfamilies of the Hystrichopsyllidae, as well as members of a number of other flea families. A great number of endemic species in the genera Palaeopsylla and Ctenophthalmus (Hystrichopsyllidae), both in the European and Asian parts of the Palaearctic, can be explained by the junction of the European and Asian continental platforms in the late Cretaceous and their subsequent isolation during the Paleocene. A considerable contribution to the flea fauna in the Russian territory was made by the East Asian-Nearctic center of taxonomic diversity, with a smaller role of the European palaeofauna. Immigration of species of the family Pulicidae from the Afrotropical Region is restricted to the southern territories of Russia.     

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     

The fauna of bloodsucking insects of Northwestern Russia. Characteristics of the ranges

The fauna of the Northwestern region of Russia comprises 285 species of bloodsucking insects. The number of species of mosquitoes and lice in the Northwest constitute 40% of the total numbers in the Russian fauna; that of the biting midges, 38%; horseflies, 35%; blackflies, 30%, and fleas, 19%. Representatives of 4 dipteran families of the “gnus” complex, and also lice and fleas of the Northwestern Russian fauna possess ranges of 12 types, most of which include large part of the Holarctic or Palaearctic; 57 species (20%) of the species have Holarctic ranges; 82 species, or 28%, trans-Palaearctic ranges; 49 species, or 17%, Western-Central Palaearctic ranges; and 73 species, or 25%, Western-Palaearctic ranges. The fraction of species (8) with other range types is 19%. A synopsis of the species groups characterized by these types of ranges is given. Species with ranges extending as far northwards as the tundra zone constitute 13%, those with ranges including the taiga zone, 60%, and those with other range types, 27% of the fauna.     

Distribution of fleas in rodent colonies in the Northwestern Precaspian region

Quantitative characteristics of phoresy of fleas on their rodent hosts inhabiting the sand landscapes of the northwestern Precaspian region were obtained. The structure of parasitic contact networks formed in the process was determined. The probability of a host change by fleas as the result of periodic feeding was assessed as high. The parameters of flea distribution were shown to be similar in the Northwestern Precaspian region and the Volgo-Ural sands. The ability of the flea Xenopsylla conformis to parasitize on social voles without significantly decreasing its abundance was revealed.     

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.     

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.     

Parasite-host relationship of flies and small mammals in the Omsk region

A generalized analysis of data on a flea fauna, range of their hosts in various natural zones, and features of parasite-host relationships between fleas and small mammals obtained in the Omsk province during long term researches in 1963-1997 is given. 35 flea species are recorded. The most mass species both on animals and in their nest is Ctenophthalmus assimilis; the numerous species are Amalareus penicilliger, Megabothros rectangulatus, M. walkeri, Peromyscopsylla silvatica, Ctenophthalmus unciatus, Palaropsylla sorecis, Doratopsilla birulai, Neopsylla pleskei, Hystrichopsylla talpae; the usual species--Ceratopsyllus garei, M. calcarifer, M. turbidus, Frontopsylla elata, Amphipsylla sibirica, A. kuznetzowi, Peromyscopsylla dasycnema, Radinopsylla integella, Catalagia dacenkoi. Other species are less numerous or infrequent. The general infection rate of the flea populations on rodents and insectivores makes 30.4%. For certain species it reaches 65.1% (on red-backed vole Cletrionomys rutilus), for regular groups--86.9% (on shrews of the genus Sorex). The greatest variety of the flea populations is observed on Microtus arvalis, the least one--on Sorex caecutiens and S. daphaenodon. In the nests of small mammals the variety of fleas is significantly lower. Based on the index of flea species relative "loyalty" to small mammals and their nest we have recognized 6 groups of fleas.     

Patterns of formation of the flea (Siphonaptera) fauna in the Caucasus

Fleas of the Caucasus belong to 155 species of 40 genera, constituting 17% and 43% of the species and generic composition of the Palaearctic fauna, respectively. The Caucasian fauna includes 23 endemic species but no endemic genera or subgenera. In the number of species, the Caucasian fauna is similar to that of the Mediterranean Subregion and is significantly poorer than the faunas of the Euro-Siberian (by 2.2 times) and Irano-Turanian (by 1.7 times) Subregions. Based on taxonomic diversity, we can propose a hypothesis on the West and East Palaearctic sources of the Caucasian fauna. The West Palaearctic source has determined the distribution of pulicomorph fleas of the families Pulicidae and Coptopsyllidae from Africa, on the one hand, and of fleas of the genera Ctenopthalmus and Palaeopsylla from Europe, on the other hand. Fleas of the Holarctic genera, such as Ceratophyllus and Megabothris, entered the Caucasus by the north Asian route; fleas of the genera Neopsylla, Rhadinopsylla, and Hystrichopsylla migrated to the Caucasus from east and central Asia by the south Asian route, through Middle and Western Asia.     

Nested pattern in flea assemblages across the host's geographic range

Understanding non-random patterns in the taxonomic composition of communities occurring in insular or fragmented habitats remains a major goal of ecology. Nested subset patterns are one possible departure from random community assembly that has been reported for communities of both free-living and parasitic animals. Here, we investigate the effects of extrinsic factors on the occurrence of nestedness among the assemblages of fleas found in different populations of the same host species, using data on 25 mammalian host species. The patterns of flea species composition among host populations spanned the entire spectrum from significantly nested to significantly anti-nested. After controlling for host phylogeny, we found that across host species, the tendency for flea assemblages to approach nestedness increased with increasing host geographic range size and with decreasing latitude of the host's geographic range. This tendency also decreased with an increase in a composite variable combining data on mean January and July temperature. The number of closely-related mammalian species living in sympatry with a given host species had no influence on whether or not the structure of flea assemblages among its populations departed from randomness. We propose explanations for these results that include: the possible gradual loss of flea species as a host expands its range from its initial area of origin, the lack of specific flea faunas in narrowly-distributed host species, interspecific differences in the dispersal abilities of flea species becoming amplified in hosts with broad geographical ranges, and the effect of latitude, climate and environment on the probabilities of host-switching and extinction in fleas. Overall, our results suggest that the structure of flea assemblages in mammalian hosts may be driven by features of host biology.     

Abundance patterns of two Oropsylla (Ceratophyllidae: Siphonaptera) species on black-tailed prairie dog (Cynomys ludovicianus) hosts.

Behavioral, genetic, and immune variation within a host population may lead to aggregation of parasites whereby a small proportion of hosts harbor a majority of parasites. In situations where two or more parasite species infect the same host population there is the potential for interaction among parasites that could potentially influence patterns of aggregation through either competition or facilitation. We studied the occurrence and abundance patterns of two congeneric flea species on black-tailed prairie dog (Cynomys ludovicianus) hosts to test for interactions among parasite species. We live-trapped prairie dogs on ten sites in Boulder County, CO and collected their fleas. We found a non-random, positive association between the two flea species, Oropsylla hirsuta and O. tuberculata cynomuris; hosts with high loads of one flea species had high loads of the second species. This result suggests that there is no interspecific competition among fleas on prairie dog hosts. Host weight had a weak negative relationship to flea load and host sex did not influence flea load, though there were slight differences in flea prevalence and abundance between male and female C. ludovicianus. While genetic and behavioral variation among hosts may predispose certain individuals to infection, our results indicate apparent facilitation among flea species that may result from immune suppression or other flea-mediated factors.     

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.     

Taxonomic composition and zoogeographic characteristics of the flea fauna (Siphonaptera) of Russia

255 species and 59 subspecies of fleas from 55 genera of 7 families are known from Russia, which is 30% of the Palaearctic fauna. Additionally, over 187 species of 47 genera from 7 families are known from the neighboring territories of Central and Southern Europe, Transcaucasia, Kazakhstan, Middle Asia, Mongolia, Northeast China, and Japan. 13 species of 12 genera are known only from Russia. Noteworthy is the low percent of endemic species (not more than 4%) and genera (one genus) in the Russian fauna. The principal centers of taxonomic diversity in the Palaearctic, including many endemic species and genera, lie in the Eastern Asian, Central Asian, and Turano-Iranian Subregions, outside Russia and the Euro-Siberian Subregion. The bulk of the Russian fauna is formed by the species and genera of the three largest flea families: Hystrichopsyllidae, Ceratophyllidae, and Leptopsyllidae. The family Ceratophyllidae has the greatest number of genera in the Russian fauna, and Hystrichopsyllidae, the greatest number of species. Western (Western and Western-Central Palaearctic; 84 species from 41 genera of 7 families) and Eastern (Central-Eastern and Eastern Palaearctic; 78 species from 42 genera of 6 families) species are nearly equally represented in the Russian fauna.     

Ways of the fleas (Siphonaptera) fauna formation in the Caucasus

Fleas fauna of the Caucasus is considered, possible ways of its formation are discussed. Caucasian fleas belong to 155 species and 40 genera; 23 species are endemics. Hypothesis on Western Palearctic and Eastern Palearctic sources of the Caucasian fleas' fauna formation are proposed.     

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.     

Peculiarities of thoracic and abdominal combs of fleas (Siphonaptera)

The structure of pseudosetae, spinelets, and spines of combs (ctenidia) was studied by means of light and SE microscopy in 80% of genera and subgenera of the World fauna. It is found out that peculiarities of ctenidiae in the prothorax and in tergites of the abdomen are characteristics of families and infraorders of fleas. Some characters of ctenidiae found in certain flea genera are reductions and apparently caused by habitation in some extremal conditions. An absence of ctenidiae in the unfraorder Pulicomorpha is compensated by more developed posterior margin of prothorax and general abbreviation of all thoracal segments. Reasons of ctenidiae absence, which is observed in certain genera of the infraorders Ceratophyllomorpha, Pygiopsyllomorpha and Hystricopsillomorpha associated with the same hosts, is not clear. It is confirmed, that distance between ctenidiae in different flea species associated with the same species host species, however it is recovered, that this distance correlates with the diameter of most thin hair of host. In some flea species the distance between ctenidia spices in females is larger, than in males. It is found, that sexual dimorphism by this character may not be expressed in certain species of closely related species group of fleas. It is suggested that ctenidiae were present even in the common ancestor of fleas. The hypothesis on origin of spines and pseudosetae from setae of the posterior walls of toracal and abdominal segments in the common ancestor of fleas is proposed.     

Of fleas and geese: the impact of an increasing nest ectoparasite on reproductive success

Past studies on the relationship between nest ectoparasites and avian fitness have been primarily limited to altricial hosts. Life history strategies of precocial and altricial birds vary considerably, limiting our ability to infer the effect of nest parasites on fitness of precocial species. Ross's Chen rossii and lesser snow goose Chen caerulescens caerulescens populations have been growing at unprecedented high rates. New limiting factors on vital rates of these precocial birds may arise after populations have been released from previously regulating factors. The flea Ceratophyllus vagabundus vagabundus is an apparently newly emerging nest parasite in the arctic goose colony at Karrak Lake, Nunavut, Canada. We examined the relationship between flea abundance (measured by the proportion of goose eggs covered by blood in each nest) and goose reproductive success from 2001–2004. In three of four years of study, nest success was inversely related to flea abundance in nests. Despite the potential for high costs to individuals, the overall effects of fleas on goose nesting success have thus far been small. We demonstrated that nest parasites negatively influence reproductive success of precocial bird hosts despite host life history strategy of leaving the nest quickly after hatch, which results in minimal exposure to nest parasites compared to altricial birds that raise their young in the nest.     

Survey of Flea Infestation in Dogs in Different Geographical Regions of Iran

Medically important arthropods, including fleas, play an important role in causing clinical disorders and disease in man and domestic animals. This study was conducted to determine the seasonal flea infestations for domestic dogs from different geographic regions of Iran. A total of 407 fleas, belonging to 5 different species, were recovered from 83 domestic dogs from 3 regions. There was a distinctive pattern of species distribution and infestations with the highest infestation rates observed in a temperate climate and higher rainfall. Additionally, fleas were observed over all seasons, except February and March, with the highest infestation rate observed in August (24.7%) and the lowest rate in January (1.7%). They also parasitize dogs with a different spectrum of species. The cat flea, Ctenocephalides felis (67.5%), exhibited the highest prevalence among all flea species found on dogs. Thus, climatic conditions and seasonal patterns impact on flea infestation and must be considered in developing control programs.     

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.