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I S Khokhlova

Publications and source records attributed to I S Khokhlova.

17 recordsLinked to original sources

Age, intensity of infestation by flea parasites and body mass loss in a rodent host.

Parasitism by the flea Synosternus cleopatrae does not affect the body mass of its principal rodent host, Gerbillus andersoni under natural infestation levels. We hypothesized that the lack of negative effects of flea parasitism on rodent body mass could be related either to the low level of natural infestation or to the differential susceptibility of rodent age cohorts to flea parasitism. We tested these hypotheses by measuring body mass change under flea parasitism in (a) adult rodents infested with fleas above the natural infestation level (the first hypothesis) and (b) juvenile rodents infested with fleas at natural infestation levels (the second hypothesis). Adult individuals parasitized by a number of fleas higher than in nature lost body mass at higher rates than non-parasitized control individuals. Parasitism significantly affected daily body mass change of juvenile gerbils. Juvenile rodents parasitized by fleas at the natural level of infestation lost body mass faster and gained body mass slower than control animals. We suggest that some regulating mechanisms may limit natural flea densities at a point at which the negative effect on hosts is below the accuracy of our measurements. However, natural flea densities are sufficiently high to harm the more susceptible, juvenile cohort.

Age Factors↗

Resource predictability and host specificity in fleas: the effect of host body mass.

Ecological specialization is hypothesized to result from the exploitation of predictable resource bases. For parasitic organisms, one prediction is that parasites of large-bodied host species, which tend to be long-lived, should specialize on these hosts, whereas parasites of small host species, which represent more ephemeral and less predictable resources, should become generalists. We tested this prediction by quantifying the association between the level of host specificity of fleas and the mean body mass of their mammalian hosts, using published data from 2 large, distinct geographical regions (South Africa and northern North America). In general, we found supporting evidence that flea host specificity, measured either as the number of host species exploited or their taxonomic distinctness, became more pronounced with increasing host body mass. There were, however, some discrepancies among the results depending on the different measures of host specificity, the geographical region studied, or whether we used the raw values or phylogenetically independent contrasts. These are discussed with respect to other forces acting on the evolution of host specificity in parasites, as well as in the context of the regions' contrasting evolutionary histories. Overall, though, our findings indicate that the exploitation of large-bodied, and therefore long-lived, host species has promoted specialization in fleas, most likely because these hosts represent predictable resources.

Animals↗

Covariance in species diversity and facilitation among non-interactive parasite taxa: all against the host.

Different parasite taxa exploit different host resources and are often unlikely to interact directly. It is unclear, however, whether the diversity of any given parasite taxon is indirectly influenced by that of other parasite taxa on the same host. Some components of host immune defences may operate simultaneously against all kinds of parasites, whereas investment by the host in specific defences against one type of parasite may come at the expense of defence against other parasites. We investigated the relationships between the species diversity of 4 higher taxa of ectoparasites (fleas, sucking lice, mesostigmatid mites, and ixodid ticks), and between the species richness of ectoparasites and endoparasitic helminths, across different species of rodent hosts. Our analyses used 2 measures of species diversity, species richness and taxonomic distinctness, and controlled for the potentially confounding effects of sampling effort and phylogenetic relationships among host species. We found positive pairwise correlations between the species richness of fleas, mites and ticks; however, there was no association between species richness of any of these 3 groups and that of lice. We also found a strong positive relationship between the taxonomic distinctness of ecto- and endoparasite assemblages across host species. These results suggest the existence of a process of apparent facilitation among unrelated taxa in the organization of parasite communities. We propose explanations based on host immune responses, involving acquired cross-resistance to infection and interspecific variation in immunocompetence among hosts, to account for these patterns.

Animals↗

Abundance and distribution of fleas on desert rodents: linking Taylor's power law to ecological specialization and epidemiology.

We investigated variation in the abundance-prevalence relationships of fleas among 17 different flea-host associations as well as among different species of hosts and fleas in the Negev desert. We explored variation in the value of exponent of Taylor's power relationship with changes in flea community size and flea specialization (host specificity and seasonal pattern of activity). We tested if a simple epidemiological model can reproduce the pattern of the abundance-prevalence relationship. We confirmed aggregated distribution of fleas within a population of host species as well as across a whole host community and the existence of a positive relationship between local flea abundance and their prevalence. Prevalence, mean abundance and variance of abundance were significantly higher in host specific than host opportunistic fleas. When ecological specialization was considered, based on a seasonal pattern of activity, these parameters were higher in year-round-active than seasonal fleas. The degree of flea specialization and flea community richness affected the pattern of the relationship between mean abundance and its variance. Power law slopes decreased with increasing richness of flea community. A simple epidemiological model based on mean flea abundance and degree of aggregation, corrected for host sample size, can predict the observed pattern of prevalence. In some cases, observed flea prevalence was higher than that predicted from the epidemiological model. The discrepancy of the observed prevalence from that predicted by the model can be explained by either a relatively low negative effect of flea parasitism on a host (at least, in terms of pathology) or strong resistance of a host to flea parasitism or both.

Animals↗

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.

Age Factors↗

Relationships between parasite abundance and the taxonomic distance among a parasite's host species: an example with fleas parasitic on small mammals.

Opportunistic parasite species, capable of exploiting several different host species, do not achieve the same abundance on all these hosts. Parasites achieve maximum abundance on their principal host species, and lower abundances on their auxiliary host species. Taxonomic relatedness between the principal and auxiliary host species may determine what abundance a parasite can achieve on its auxiliary hosts, as relatedness should reflect similarities among host species in ecological, physiological and/or immunological characters. We tested this hypothesis with fleas (Siphonaptera) parasitic on small Holarctic mammals. We determined whether the abundance of a flea in its auxiliary hosts decreases with increasing taxonomic distance of these hosts from the principal host. Using data on 106 flea species from 23 regions, for a total of 194 flea-locality combinations, we found consistent support for this relationship, both within and across regions, and even after controlling for the potentially confounding effect of flea phylogeny. These results are most likely explained by a decrease in the efficiency of the parasite's evasive mechanisms against the host's behavioural and immune defences with increasing taxonomic distance from the principal host. Our findings suggest that host switching over evolutionary time may be severely constrained by the coupling of parasite success with the relatedness between new hosts and the original host.

Animals↗

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.

Animals↗

Host specificity and foraging efficiency in blood-sucking parasite: feeding patterns of the flea Parapulex chephrenis on two species of desert rodents.

Parasite species can adapt to ecological, behavioral, physiological and biochemical traits of a particular host species. The flea Parapulex chephrenis occurs on the spiny mouse Acomys cahirinus, but does not occur on a co-existing gerbil, Gerbillus dasyurus. To test the hypothesis that the host species affects feeding parameters of a host-specific flea, we studied the feeding rate, rate of blood digestion and resistance to starvation of P. chephrenis when feeding on A. cahirinus and G. dasyurus. We predicted that P. chephrenis would: (1) fill its gut with blood faster, (2) digest blood for a shorter time, and (3) survive longer when starved while feeding on its specific host, A. cahirinus, than on a non-specific host, G. dasyurus. These three responses were observed when P. chephrenis fed on the different hosts and, consequently, our predictions were supported. Twenty percent of fleas filled their midgut after feeding for 10 min on A. cahirinus but this occurred only after 25 min on G. dasyurus. The middle stage of blood digestion was significantly shorter in all fleas feeding on A. cahirinus than in fleas feeding on G. dasyurus. Flea survival was shorter when feeding on G. dasyurus than when feeding on A. cahirinus at 25 degrees C, but no difference in survival time was found at 15 or 20 degrees C. Both A. cahirinus, the specific host, and G. dasyurus, the non-specific host, co-exist in rocky habitats, yet P. chephrenis occurs on one rodent and not the other. The absence of P. chephrenis on G. dasyurus in nature and the decreased foraging efficiency when feeding on this species in the laboratory suggests that some physiological and biochemical differences between hosts can lead to sharp ecological differences in host-parasite relationships.

Animals↗

Annual cycles of four flea species in the central Negev desert.

Bionomics of fleas (Siphonaptera) parasitizing rodent hosts, mostly the gerbil Gerbillus dasyurus (Wagner) and the jird Meriones crassus Sundevall (Gerbillidae), were investigated in the central Negev desert of Israel. Populations were sampled weekly (by Sherman trapping of hosts) from August 2000 to July 2001. Among 1055 fleas of nine species captured, four species predominated (94%). Two species of Pulicidae, Xenopsylla dipodilli Smit and X. ramesis (Rothschild), reproduced perennially, whereas adults of Nosopsyllus iranus theodori Smit (Ceratophyllidae) and Stenoponia tripectinata medialis Jordan (Hystrichopsyllidae) occurred only in cool months (October-March). During their main activity season on the most infested host species (estimated from > 300 trap-nights/month), prevalence of these four flea species reached 40-70%, 20-30%, 100% and 50-70%, respectively, with infestation intensities of 2-2.7, 7-12, 2-3.5 and 2.5-7 fleas per infested rodent, respectively. Xenopsylla dipodilli oviposition peaked during autumn with parous rate > 80% by September-October. During December-April, the majority of X. dipodilli females were immature and/or nulliparous (defined as having mature ovaries but no follicular relics). In contrast, X. ramesis had two reproductive peaks, in mid-spring and autumn, evidenced by the influx of immature females in late spring and summer (30-40%) and in winter (20-30%) after maximal parous rates: 80-100% in March-April and 95-100% in October-November. Nosopsyllus iranus theodori and Stenoponia tripectinata medialis adults occurred only during cool months. At the beginning of activity, during October-November, the sex ratio of N. i. theodori was strongly biased to females (86%) that were immature and/or nulliparous. In winter, adult females were 52-65% parous and 10-32% immature. In March, as the adult population of N. i. theodori declined, 78% of females were parous and 12% immature. Seasonal activity of S. t. medialis (November-March) was shorter than for the other three species; females were predominantly nulliparous in November (80%), after which the proportion of parous females increased gradually to 84% in February. Two females of S. t. medialis collected in March were mature but nulliparous, suggesting that this species of flea might 'oversummer' (as pupae or teneral adults) in the cocooned stage.

Animals↗

The effect of substrate on survival and development of two species of desert fleas (Siphonaptera: Pulicidae).

Fleas Xenopsylla conformis mycerini and Xenopsylla ramesis replace each other on the same rodent host (Meriones crassus) in two habitats that differ in substrate texture (sand and loess-like sediments, respectively). We hypothesized that the substrate is an important factor determining flea distribution and studied survival of larvae, pupae and newly emerged adults as well as the rate of pre-imaginal development of these flea species in sand and loess rearing medium (= substrate). Texture of rearing medium did not affect survival and development rate of eggs in either X. c. mycerini or X. ramesis. Larval survival and the rate of development were both affected by the factor of substrate. Survival of X. c. mycerini larvae was significantly higher in sand than in loess substrate, whereas survival of X. ramesis larvae did not differ in different substrates. Larvae of both species developed faster in sand substrate than in loess substrate. Maximal survival time of X. c. mycerini larvae that died before pupation did not depend on substrate, whereas X. ramesis larvae survived significantly longer in loess than in sand substrate. Most pupae of both species survived successfully on both substrates, but the duration of pupal stages in sand substrate was longer than that in loess substrate in both species. Newly emerged adults of both species survived similar time in both sand and loess substrate. Irrespective of substrate, adult X. c. mycerini survived for a shorter time than did adult X. ramesis. No between-sex within-species differences in survival time of newly emerged adults in sand versus loess substrate were found in X. c. mycerini. Survival time of males and females of X. ramesis differed in sand substrate but not in loess substrate.

Animals↗

Time of survival under starvation in two flea species (Siphonaptera: Pulicidae) at different air temperatures and relative humidities.

We studied the effect of air temperature and RH on the survival time of adult Xenopsylla conformis Wagner, 1903 and Xenopsylla ramesis Rothschild, 1904 fleas during starvation to explain the paratopic habitat distribution of these species on opposite ends of a precipitation and temperature gradient in the Negev Highlands, Israel. We hypothesized that the pattern of distribution of the two flea species is caused by differential microclimatic preferences of imagoes and predicted that (1) the resistance to starvation would differ between the two flea species at different air temperatures (10 degrees C - 38 degrees C) and relative humidities (RH) (40%-92%) and (2) survival time of starving X conformis would be longer than that of starving X ramesis at high air temperatures and low RHs. Survival time of newly emerged X conformis was dependent on air temperature but not on RH, whereas in newly emerged X ramesis it was affected by both air temperature and RH. Generally, survival time of newly emerged fleas was longer at lower air temperatures and higher humidities than at higher air temperatures and lower humidities. No significant difference in survival time between species in dependence on either air temperature or RH were found for newly emerged fleas. Fed fleas of both species responded similarly to changes in air temperature and RH in terms of survival time. Survival time at lower temperatures was longer than that at higher temperatures. Females survived longer than males at all air temperatures except for the highest temperature when the survival time of both sexes was similarly low. In both species, the effect of RH on survival time was significant at the highest RH only, with a decrease in survival time at 92% RH. In contrast, survival times at lower RHs did not differ. Males of both species survived for less time than females at all RHs. X conformis generally survived for less time than X ramesis, all else being equal. The only regime at which X conformis survived longer than X. ramesis was 38 degrees C and 40% RH. Newly emerged fleas survived for significantly longer time than fed fleas.

Adaptation, Physiological↗

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.

Animals↗

Effect of air temperature and humidity on the survival of pre-imaginal stages of two flea species (Siphonaptera: Pulicidae).

The survival of immatture fleas at 25 and 28 degrees C and 40, 55, 75, and 92% RH was studied to test the hypothesis that the difference in microclimatic preferences determines habitat distribution of Xenopsylla conformis Wagner, 1903 and Xenopsylla ramesis Rothschild, 1904. Survival of X. conformis eggs did not depend on either temperature or humidity or both, whereas eggs of X. ramesis survived significantly less at 40% RH than at higher humidities. No larva of either species survived at 40% RH at either temperature. Larval survival of both species at both temperature regimes was significantly lower at 55% humidity than at higher humidities. Maximal survival time of larvae that died before pupation depended on both temperature and humidity in both species. Change of humidity during early stages of the life cycle (from egg to larva) increased the maximal survival time in X. conformis larvae but decreased that in X. ramesis larvae. Pupal survival was higher at higher humidities independent of temperature. Survival of X. conformis pupae was lower than that of X. ramesis pupae when the relative humidity was low. Humidity change on later stages (from larva to cocoon) decreased X. conformis pupal survival and had no effect on X. ramesis pupal survival. The sex ratio of emerged adults was not affected by either temperature or humidity in both species. Changes in humidity between egg and larval emvironments significantly decreased the percentage of females in X. conformis emergence at 28 degrees C.

Air↗

The effect of behavioural interactions on the transfer of fleas (Siphonaptera) between two rodent species.

The transfer of fleas between the house mouse, Mus musculus, and the midday jird, Meriones meridianus, through social contacts was studied by performing interspecific dyadic encounters on a neutral arena. M. meridianus and M. musculus exchanged parasites through social contacts. The intensity of this exchange was higher between mice and adult male and young jirds than between mice and adult female jirds. The pattern of behavioural interactions was the main determinant of the intensity of flea exchange. Adult female jirds were highly aggressive towards mice. In contrast, interactions between male and young jirds and mice were mainly cohesive and resulted in higher frequency of flea transfer. The proportion of fleas transferred was positively correlated with the number and cumulative duration of tactile contacts between opponents in dyads.

Aggression↗

Host-habitat relations as an important determinant of spatial distribution of flea assemblages (Siphonaptera) on rodents in the Negev Desert.

We studied flea assemblages on rodents in different habitats of the Ramon erosion cirque in the Negev Desert to examine whether host-habitat relations influence flea spatial distribution. Eleven flea species parasitizing 12 rodent species were recorded. There was significant positive relationship between flea species richness and body mass of the host species; no relationships were found between relative richness of flea assemblage and either the number of habitats occupied by the host species or the size of host geographical range. The differences in pattern of flea parasitism among habitat types within host species were determined by both environmental features of a habitat and the specific pattern of habitat use by rodents. There was replacement of Xenopsylla conformis by Xenopsylla ramesis on Meriones crassus and Gerbillus dasyurus among different habitats. The results of ordination of the flea collections from each individual host demonstrated that the flea assemblages were segregated mainly along 4 axes, which explained 86% of total variance. Each of the ordination axes corresponded with a change in flea species composition. The directions of these changes were (1) among-hosts within a habitat and (2) among-habitats within a host.

Animals↗

Level of energy intake affects the estrous cycle in Sundevall's jird (Meriones crassus).

Effects of energy intake on the estrous cycle of the desert gerbillid, Sundevall's jird (Meriones crassus; 80 g; n=22) were studied. Females were offered either maintenance or below maintenance levels of millet seeds and ad lib. Atriplex halimus leaves and stems; drinking water was not available. Vaginal smears were used to determine sexual stage. We hypothesized that the estrous cycle ceases at low levels of energy intake and commences when sufficient energy is available. Females lost body mass linearly with a decrease in metabolizable energy intake. Estrous cycle averaged 4.46 d at maintenance energy intake but increased to an average of 7.81 d at 70% of maintenance energy intake. A cessation of the cycle occurred at an energy intake below 70% of maintenance requirements, which resulted in a body mass loss of more than 1% per day. More variability in the length of the different stages of the cycle was found with lower levels of energy intake. When offered ad lib. millet seeds and A. halimus (n=14), recovery to the normal cycle was attained within 10 d by 43% of the females and within 16 d by the rest of the females. Recovery time was longest in females that previously had the lowest energy intake. We concluded that the estrous cycle of M. crassus is sensitive to energy intake. With restricted energy intake, the estrous cycle and reproductive activities cease but can be restored with provision of adequate energy. This strategy ensures that reproduction occurs when conditions of food availability and body condition of the females are favorable.

Animals↗

Fiber digestion and energy utilization of fat sand rats (Psammomys obesus) consuming the chenopod Anabasis articulata.

Most herbivorous rodents consume a variety of plants and, when available, select ones low in fiber content. In contrast, the fat sand rat (Psammomys obesus), a diurnal gerbillid rodent that is wholly herbivorous, is able to survive while consuming only the halophytic chenopod Anabasis articulata. We hypothesized that these gerbils are able to digest chenopods efficiently and that fiber digestion contributes substantially to their energy budget. Digestibility of total fibers (NDF), hemicellulose, and cellulose was 51.6%, 66.4%, and 44.0%, respectively. Energy derived from fiber digestion was 211.7 kJ kg(-0.75) d(-1) and resulted mostly from hemicellulose digestion. Average daily metabolic rate was 658.4 kJ kg(-0.75) d(-1), and basal metabolic rate (BMR) was determined to be close to 192.6 kJ kg(-0.75) d(-1). Therefore, fiber digestion provided 32% of maintenance requirements and 110% of BMR requirements, one of the highest values reported thus far for placental mammals. The efficiency of utilization of the chenopod was 0.29, a low value compared to other dietary items. We concluded that A. articulata fulfills all the energy and nutrient requirements of fat sand rats, even though energy yields and efficiency of utilization of its energy is low.

Adaptation, Physiological↗