Morphological diversity of the skeletal structures of fleas (Siphonaptera). Part 2: The general characteristic and features of the thorax

Analysis of a considerable amount of data on the anatomy of 27 structures of the thorax in 96 genera of fleas (over 90% of the genera in the world fauna) shows that different flea taxa can be described based on 48 universal and specific characters, whose 203 states reflect the entire known diversity of the flea morphology. Of them, 26 characters with 96 states can be formulated based on universal terms; 22 universal characters with 88 states describe the proportions of the segments and their parts, and also proportions and shapes of their sclerites; 23 specific characters with 107 states describe the structure of the thoracic sclerites and ridges as well as the patterns of their junctions. Judging by the number of characters and their states, the most evolutionarily flexible structures in fleas are the mesosternite and its apodeme together with the upper part of the mesopleural rod. Homoplasies at various levels comprise less than half (40%) of all the character states in the thorax structure. The character states reflecting the phylogenetic closeness of taxa make up about 15%.     

Morphological diversity of the skeletal structures of fleas (Siphonaptera). Part 3: The general characteristics and features of thoracic setation

The paper continues the series of publications (Medvedev, 2015, 2016) devoted to the analysis of the structural features of fleas. In this part, the structural features of the thoracic setation are analyzed for the first time. Six characters with 53 states were distinguished in the chaetome of the pronotum, mesothorax, and metathorax. The analysis reveals some cases of the formation of externally similar states due to reduction or, by contrast, strengthening of the chaetome in some genera belonging to different superfamilies, in relation to the specific traits of parasitism. In spite of high variation in the number and arrangement of thoracic setae, some character states can be used as markers of flea taxa of the family and superfamily rank.     

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.     

Cytogenetics of fleas (Siphonaptera: Pulicidae). 1. Rat fleas of the genus Xenopsylla

Five populations of Xenopsylla cheopis exhibit a chromosome complement of 2n = 17, X1X2Y (male), and 2n = 18, X1X1X2X2 (female). A detailed analysis of populations of X. astia from Bombay and Trivandrum led to the identification of two distinct cytotypes which hybridisation studies indicated were sibling species. These are referred to as X. astia with a diploid chromosome number of 2n = 18, X1X2X3Y (male), and 2n = 20, X1X1X2X2X3X3 (female) and X. prasadii with 2n = 10, X1X2Y1Y2 (male), and 2n = 10 X1X1X2X2 (female). It is proposed that X. prasadii is derived from X. astia through translocation/fusion events since the average total chromosome lengths are remarkably similar in all three species.     

Bird fleas (Insecta: Siphonaptera): taxonomic diversity, biogeographical distribution

If Mammals are the primary hosts of Siphonaptera, 6% of them have changed their trophic appetency for Birds. What are the reasons, what are the adaptations to be adopted by Fleas, what are the families or species groups of fleas concerned, and at last what are the host-families? As to this last question, it is clear that deviation was ecological but not phyletical.     

Morphological Diversity of the Skeletal Structures and Problems of Classification of Fleas (Siphonaptera). Part 6

The structure of 76 skeletal elements of adult fleas was analyzed, and the distribution of 114 characters with 446 character states over the body tagmata, segments, and morphofunctional complexes was investigated. Among them, 40% of the characters (40) and their states (163) describe the diversity of the structures of the frontal complex (including those of the head and prothorax), which is related to the specific features of flea parasitism. A large part of the characters (18) and their states (83) describe the structures of the nototrochanteral complex of the meso- and metathorax responsible for jumping. The total number of all types of homoplasies (258 states) is almost 1.8 times as great as the number of the states (145) that may be regarded as synapomorphies. The ancestral states (43) comprise a smaller portion of the total number. The proportion of the synapomorphic and homoplastic character states varies between the morphofunctional complexes.     

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.     

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.     

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.     

Experimental interspecific hybridization in fleas of the genus Nosopsyllus (Siphonaptera: Ceratophyllidae)]

Experimental interspecific hybridization between males and females of Nosopsyllus fasciatus and N. mokrzeckyi was carried out. Most abundant progeny was obtained after hybridization between N. fasciatus females and N. mokrzeckyi males. Hybrid descendants inherited, in general, the characters of the maternal species and in less number the paternal characters and those of both parents. The fecundity of hybrid progeny of N. fasciatus females and N. mokrzeckyi males in two generations was studied. A conclusion is made concerning close affinity of the species and their relative evolutionary youth.     

A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations

Siphonaptera (fleas) is a highly specialized order of holometabolous insects comprising ～2500 species placed in 16 families. Despite a long history of extensive work on flea classification and biology, phylogenetic relationships among fleas are virtually unknown. We present the first formal analysis of flea relationships based on a molecular matrix of four loci (18S ribosomal DNA, 28S ribosomal DNA, Cytochrome Oxidase II, and Elongation Factor 1‐alpha) for 128 flea taxa from around the world representing 16 families, 25 subfamilies, 26 tribes, and 83 flea genera with eight outgroups. Trees were reconstructed using direct optimization and maximum likelihood techniques. Our analysis supports Tungidae as the most basal flea lineage, sister group to the remainder of the extant fleas. Pygiopsyllomorpha is monophyletic, as are the constituent families Lycopsyllidae, Pygiopsyllidae, and Stivaliidae, with a sister group relationship between the latter two families. Macropsyllidae is resolved as sister group to Coptopsyllidae with moderate nodal support. Stephanociricidae is monophyletic, as are the two constituent subfamilies Stephanocircinae and Craneopsyllinae. Vermipsyllidae is placed as sister group to Jordanopsylla. Rhopalopsyllidae is monophyletic as are the two constituent subfamilies Rhopalopsyllinae and Parapsyllinae. Hystrichopsyllidae is paraphyletic with Hystrichopsyllini placed as sister to some species of Anomiopsyllini and Ctenopariini placed as sister to Carterettini. Ctenophthalmidae is grossly paraphyletic with the family broken into seven lineages dispersed on the tree. Most notably, Anomiopsyllini is paraphyletic. Pulicidae and Chimaeropsyllidae are both monophyletic and these families are sister groups. Ceratophyllomorpha is monophyletic and includes Ischnopsyllidae, Ceratophyllidae, and Leptopsyllidae. Leptopsyllidae is paraphyletic as are its constituent subfamilies Amphipsyllinae and Leptopsyllinae and the tribes Amphipsyllini and Leptopsyllini. Ischnopsyllidae is monophyletic. Ceratophyllidae is monophyletic, with a monophyletic Dactypsyllinae nested within Ceratophyllinae, rendering the latter group paraphyletic. Mapping of general host associations on our topology reveals an early association with mammals with four independent shifts to birds. © The Willi Hennig Society 2008.     

Investigation of Bartonella acquisition and transmission in Xenopsylla ramesis fleas (Siphonaptera: Pulicidae)

Bartonella are emerging and re-emerging pathogens affecting humans and a wide variety of animals including rodents. Horizontal transmission of Bartonella species by different hematophagous vectors is well acknowledged but vertical transmission (from mother to offspring) is questionable and was never explored in fleas. The aim of this study was to investigate whether the rodent flea, Xenopsylla ramesis, can acquire native Bartonella from wild rodents and transmit it transovarially. For this aim, Bartonella-free laboratory-reared X. ramesis fleas were placed on six naturally Bartonella-infected rodents and six species-matched Bartonella-negative rodents (three Meriones crassus jirds, two Gerbillus nanus gerbils and one Gerbillus dasyurus gerbil) for 7 days, 12-14h per day. The fleas that were placed on the Bartonella-positive rodents acquired four different Bartonella genotypes. Eggs and larvae laid and developed, respectively, by fleas from both rodent groups were collected daily for 7 days and molecularly screened for Bartonella. All eggs and larvae from both groups were found to be negative for Bartonella DNA. Interestingly, two of five gut voids regurgitated by Bartonella-positive fleas contained Bartonella DNA. The naturally infected rodents remained persistently infected with Bartonella for at least 89 days suggesting their capability to serve as competent reservoirs for Bartonella species. The findings in this study indicate that X. ramesis fleas can acquire several Bartonella strains from wild rodents but cannot transmit Bartonella transovarially.     

Alternative prognostication of the population of fleas of the genus Xenopsylla (Siphonaptera)

A possibility of alternative prognostication of the autumn abundance of fleas was shown by means of statistical analysis of prognostication tables made up on the basis of the distribution of informative factors: indices of the abundance of the great gerbil, which is the main host of fleas, weather conditions and characteristics of the populations of the fleas.     

A morphologic analysis of digestion in Leptopsylla segnis (Siphonaptera: Leptopsyllidae) fleas

Changes in the ultrastructure of cells of the intestinal epithelium during the digestion of one blood portion were traced in the fleas L. segnis. It is shown that alongside with the cavity digestion take place elements of intracellular digestion. Hypothetic scheme of the digestive cell functioning is given.     

The plague in Vietnam: history and inventory of collected fleas (insecta, Siphonaptera) in the inhabited zones

Ten flea species are reported in anthropic zones of Vietnam. Xenopsylla vexabilis is also here included because of it has been involved in others plague's countries. Lentistivalius klossi is the only selvatic flea known as parasite of synanthropic rats. L. klossi bispiniformis (Li and Wang, 1958), first describe from Chinese specimens, is here synonymized (syn. nov.) with the nominal subspecies.