The ontogenetic variability of the morphometric traits of 2 species of ixodid ticks in the genus Rhipicephalus]

Variability of 10 morphometric characters at all phases was investigated on laboratory cultures of R. turanicus and R. bursa. It has been shown that variability increases from phase to phase. R. turanicus nymphs of both sexes differ in the length of gnathostoma and length of the 1st tarsus. In nymphs of R. bursa sexual dimorphism manifests itself on 8 characters, in all cases sizes of organs are greater in female nymphs. Besides, engorged female nymphs are reliably greater than male ones in length and mass of the body. Mature females and males of R. bursa also reliably differ in the total body length as well as in sizes of all examined structures. Females and males of R. turanicus do not differ in body length but differ in 9 other morphometric characters. Correlation analysis of characters was carried out individually for each phase and sex. Correlation coefficient between characters are most low in larvae. In R. turanicus male and female nymphs the coefficients are close. In R. bursa female nymphs correlation coefficients are noticeably lower than in male ones. The level of independence of characters in female nymphs defines the degree of manifestation of sexual dimorphism at this phase: the closer the links between characters, the lesser the number of characters revealing sexual differences. Coefficients of correlation of characters coincide in males of both species. In females of R. turanicus they are lower than in males that determines the strengthening of sexual dimorphism at the phase of imago.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sexual dimorphism of the nymphal phase of ixodid tick species in the genus Dermacentor

In four studied Dermacentor species at preimaginal phases sexual dimorphism hardly reveals itself in body size and mass of individuals. The formation of sexual distinctions at the nymphal phase concerns different characters as in different species so in populations of one species. In D. niveus male and female nymphs differ in the length of II-III palpal joints and width of gnathosoma, in D. ushakovae in the length of scutum and its proportions, in the width of gnathosoma and hypostome and in the diameter of peritreme. The sex of D. silvarum nymphs can be identified by the width of gnathosoma, length of hypostome and diameter of peritreme. Female and male nymphs of D. marginatus from the Stavropol Territory and Armenia differ in the scutum proportions and populations from the western Pamirs in the length of scutum and gnathosoma.     

The meadow tick Dermacentor reticulatus in natural foci of tick-borne encephalitis in Udmurtia

In southern Udmurtia, where occur both Dermacentor reticulatus and Ixodes persulcatus, in the years of high abundance of small mammals feeding periods of larvae and nymphs of these ticks coincide completely. The duration of simultaneous parasitism of preimago of the above ixodid species on animals is of epidemiological significance as far as within this period the infection of young stages of D. reticulatus with the virus of tick-borne encephalitis occurs. This is confirmed by the isolation of I strain of the virus from D. reticulatus collected at the state of engorged nymphs on hosts common with I. persulcatus and moulted in the laboratory.     

The manifestation of sexual dimorphism in the preimago phases of 2 species of the genus Hyalomma (Ixodoidea, Ixodidae)

Experiments on laboratory cultures of Hyalomma asiaticum and H. anatolicum have shown that at the nymphal phase sexual dimorphism manifests itself in statistically reliable differences between general sizes and mass of the body of male and female individuals. Larger nymphs moult mostly into females, small ones mostly into males. Reliable differences have been noted in the sizes of scutum, gnathosoma and its appendages in male and female nymphs of both species. At the larval phase sexual dimorphism does not manifest itself in the sizes of individuals.     

Sexual dimorphism of the larvae and nymphs of the taiga tick--Ixodes persulcatus

Manifestations of sexual dimorphism at the preimaginal phase was studied on the laboratory culture of Ixodes persulcatus. It has been found out that larger engorged larvae produce, in general, larger nymphs and smaller larvae produce smaller nymphs. The classification of engorged nymphs into large (over 2.9 mm) and small ones (less than 2.9 mm) has made it possible (with 100% probability) to obtain females from the former and males from the latter. Differences in linear sizes of scutum, gnathosoma and its appendages in male and female nymphs were determined that has made possible the identification of sex in hungry nymphs.     

Evolutionary rates of secondary sexual and non-sexual characters among birds

The rate of evolutionary morphological change in secondary sexual characters among species has traditionally been assumed to exceed that for non-sexual characters, giving rise to a larger degree of divergence. We used a large data set of independent evolutionary events of exaggerated secondary sexual feather characters across all birds to test whether that was the case. Comparative analyses revealed that secondary sexual tail feather characters diverged more than wing feathers in females, and we also found that secondary sexual head feather characters diverged more than tarsi in males, when only including intra-order comparisons in the analyses. These results are in the predicted direction, with secondary sexual characters diverging more than ordinary morphological traits, partially supporting the general impression that secondary sexual characters are more variable among species than ordinary morphological characters. However, the degree of divergence among secondary sexual characters was generally not much larger than that among ordinary characters. Some non-significant differences in divergence between secondary sexual characters and ordinary characters could be explained by the cost-reducing function of ordinary morphological traits. There was no evidence of significant differences in divergence between sexes for secondary sexual characters, maybe because of genetic correlations in morphology between the sexes. However, male tarsi diverged more than female tarsi, and sexual selection might play a role in this difference in divergence. This revised version was published online in July 2006 with corrections to the Cover Date.     

珙桐群落种间关系的研究

用方差比率法(VR)和在2×2联列表基础上应用Jaccard指数、Dice指数和Pearson相关系数、Spearman秩相关系数对卧龙自然保护区三江珙桐群落种间关系进行了分析,结果表明珙桐群落中主要种群在总体上表现出正关联,群落中种间联结的显著度较低,仅有7对联结显著,珙桐与总状山矾呈显著负联结,与稠李呈极显著负联结。大多数种对间关联程度较低,存在极大排斥性的种对较少,暗示目前珙桐群落处于较成熟阶段。研究种间关系时,种间关联性测定指标结合相关系数分析得出的结果较好,最后对珙桐的保护提出了一些建议。     

Inter- and intraspecific variation in defensive compounds produced by five neotropical stink bug species (Hemiptera: Pentatomidae)

The differences in composition of defensive secretions between nymphs, adult males and adult females of Chinavia impicticornis (=Acrosternum impicticorne), Chinavia ubica (=Acrosternum ubicum), Euschistus heros, Dichelops melacanthus and Piezodorus guildinii (Hemiptera, Pentatomidae) were analysed within and between species using compositional log-ratio statistics and canonical variates analysis. Differences in composition between nymphs, males and females were found for all species, as well as when all species were pooled. In particular, tetradecanal appears to be a predominantly nymphal compound in D. melacanthus, E. heros and P. guildinii. In the two Chinavia species 4-oxo-(E)-2-hexenal and an unknown compound were more dominant in nymphs. The interspecific analysis revealed a good separation of defensive compounds according to their taxonomic relationship. Thus, the two Chinavia species grouped together, with (E)-2-decenal and (E)-2-hexenyl acetate, contributing to this separation. The other three species also differed from each other, with (E)-2-octenal associated to D. melacanthus, (E)-2-hexenal to P. guildinii and (E,E)-2,4-decadienal and tetradecanal to E. heros. The pooled analysis of stage ignoring species revealed tetradecanal and 4-oxo-(E)-2-decenal (tentative identification) strongly associated to nymphs. Thus, there are predictable differences between stages, and many of the differences are conserved between species. Consideration of these differences could prove to be important in understanding stink bug-natural enemy interactions, and in optimising biocontrol efforts.     

A morphometric study of the ant species,Myrmecia dispar (Clark) (Hymenoptera: Formicidæ)

Summary Myrmecia dispar (Clark) workers and queens show monomorphic polymorphism with limited size variation. Significant differences were found for the mean values of certain characters of workers between some colonies. Frequency histograms of head width against the number of workers in a colony varied considerably, with little or no definite pattern for the smaller colonies and ranging from unimodal to bimodal histograms for the larger colonies. Highly significant, positive, correlation values were obtained for all measured characters in each colony. Partial correlation analysis revealed much of this correlation to be due to similar growth coefficients. Regression analysis indicated that the characters head width and pronotal width were linearly related.This paper was prepared in part fulfillment of the requirements for the degree of Master of Science at the University of Sydney.     

Some aspects of intraspecific variability of the closely related species of the Dermacentor marginatus complex (Acari: Ixodidae) as demonstration of microevolutionary process

Individual and geographical variability of male conscutum pattern in five closely related species of the Dermacentor marginatus complex (D. marginatus, D. ushakovae, D. niveus, D. silvarum, D. nuttalli) has been examined. The pattern has been examined in three to six geographical samples within each specific distribution range. It was established that the pattern of the conscutum central area is subjected to individual and geographical variability on an extreme degree and gives very few intraspecific differential characters, but it shows interspecific differences in some cases. However the dark patches of marginal flank (npkappadelta and 3deltakappa respectively, see Fig. 3) show statistically significant differences in length in all geographical samples of the first four species. Characteristic features of ixodid ticks morpogenesis rule out to a considerable extent the possibility to trace geographical variability of the same characters by all stages of ontogenesis. In these cases, when it is possible, the tendencies to variability on different stages do not always coincide. There is no coordination within the species in the degree of differences of examined geographical samples by pattern and morphological characters, i. e. very complex polymorphis is observed. In the context of the evolution that kind of polymorphism of the geographical samples has provided the species with high adaptive potential for conquest of the extensive range and maintaining large numbers. The question of allopatric speciation of the closely related D. marginatus-D. silvarum with small secondary area of transgression their ranges, and sympatric speciation of the D. ushakovae-D. niveus is postulated.     

The taxonomic aspects of the transmission of the causative agent of Lyme disease

According to world literature data 17 species of ixodid ticks have been studied for natural infection with the Lyme disease agent. Analysis of the data on the level of the infection, transovarial and transphase transmission has shown that main biological vectors of Borrelia burgdorferi are the species of the subgenus Ixodes s. str. - I. ricinus, I. persulcatus (Eurasia), I. dammini, I. pacificus (North America). Potential vectors are I. scapularis, I. dentatus, Amblyomma americanum, Dermacentor variabilis. Single isolations were registered for I. neotomae, Haemaphysalis leporispalustris, D. occidentalis. Nonidentified spirocheta was isolated from A. americanum, D. variabilis, D. parumapertum, Rhipicephalus sanguineus. No agent was isolated from I. cookey, D. albipictus, R. reticulatus, H. concinna. On the basis of comparative and ontogenetic data the species from a group of main vectors: I. ricinus, I. persulcatus, I. pacificus had been attributed by me to the phyletic group persulcatus before Lyme disease was discovered and its causative agent isolated. The question whether I. scapularis belongs to the group persulcatus was also discussed at that time but left open due to somewhat aberrant structure of gnathosoma at preimaginal phases (Filippova, 1969, 1971, 1973). 6 Palaearctic, 2 Indomalayan and 3 Nearctic species were referred to the group persulcatus at the time. I. dammini was described later, in 1979. Gnathosoma of its preimaginal phases has an intermediate structure between I. scapularis and other species of the group persulcatus. Sexually mature phase and nymph of I. dentatus have much in common with Palaearctic members of the group, I. pavlovskyi, I. kazakstani, I. kashmicus. Preimaginal phases of I. scapularis and nymph of I. dentatus were studied by me on the collection material. Thus, it is possible to speak of the belonging of main vectors of B. burgdorferi to a common phyletic group within the subgenus Ixodes s. str. and, therefore, of common origin of ecological medium for the agent. At the same time each species of the vector is an evolutionally developed difference of ecological medium for B. burgdorferi. Roots of the group persulcatus could originate as far as in Paleocene before the land connection between North America and Europe disappeared. Conditions for the existence of recent species, however, appeared considerably later and their flourishing is dated by Pliocene. The main epidemiological role belongs now to I. ricinus, I. persulcatus, I. dammini, I. pacificus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Sexual size dimorphism within species increases with body size in insects

Studies examining interspecific differences in sexual size dimorphism (SSD) typically assume that the degree of sexual differences in body size is invariable within species. This work was conducted to assess validity of this assumption. As a result of a systematic literature survey, datasets for 158 insect species were retrieved. Each dataset contained adult or pupal weights of males and females for two or more different subsets, typically originating from different conditions during immature development. For each species, an analysis was conducted to examine dependence of SSD on body size, the latter variable being used as a proxy of environmental quality. A considerable variation in SSD was revealed at the intraspecific level in insects. The results suggest that environmental conditions may strongly affect the degree, though not the direction of SSD within species. In most species, female size appeared to be more sensitive to environmental conditions than male size: with conditions improving, there was a larger relative increase in female than male size. As a consequence, sexual differences in size were shown to increase with increasing body size in species with female-biased SSD (females were the larger sex in more than 80% of the species examined). The results were consistent across different insect orders and ecological subdivisions. Mechanisms leading to intraspecific variation in SSD are discussed. This study underlines the need to consider intraspecific variation in SSD in comparative studies.     

Plant smells---important determinants in the behavior and rate of development of ixodid disease vectors]

Experiments with nymphs of Ixodes persulcatus and Dermacentor marginatus have shown that the rate and degree of engorgement, dropping off from the mouse, metamorphosis longevity and weight of emerging imagoes change under plant odour influence. The influence of acetone extract vapours of pepper, poplar buds, linden and aspen was tested. Under the influence of the vapour, acting as an attractant for the adults, engorgement weight of their nymphs and the weight of imagoes emerging from them increased; attractants change the nymph-imago metamorphosis duration as well. Repellent acting odours, as a rule, produce an opposite reaction. The phenomenon of the inversion of the infected D. marginatus reaction to the odour is proved in nymphs. Attractant for naive adult ticks (poplar buds) increased the time of feeding and weight of nymphs in comparison with the control. Just the opposite: nymphs infected with tick-borne encephalitis (TBE) virus produce an opposite reaction--weight decrease under influence of poplar buds as the repellent. Importance of the above phenomena for the ticks distribution and TBE epizootiology is discussed.     

Factors influencing the degree of sexual size dimorphism within and among calanoid copepod species

Populations of Diaptomus leptopus (Copepoda: Calanoida) and other calanoid copepods exhibit varying degrees of sexual size dimorphism. We examined whether intraspecific or interspecific variation in dimorphism could be explained by allometry, and we examined the relationship between adult size attained and development rate to determine any relationship between the two. We compared the degree of sexual size dimorphism in D. leptopus and in other calanoid copepods inhabiting temporary and permanent habitats. Allometry did not explain variation in sexual size dimorphism within or among populations or among species. Permanence of habitat affected the degree of dimorphism: dimorphism was greater within and among species inhabiting temporary environments. Non-significant differences in development rate were found among populations and significant differences were found between sexes of D. leptopus when reared under identical laboratory conditions: males developed more rapidly than females but there was no general relationship between development rate and adult size. Potential adaptive hypotheses to explain the differences between populations inhabiting temporary and permanent habitats are discussed.     

Genetics of sexual isolation based on courtship song between two sympatric species: Drosophila ananassae and D. pallidosa

Sexual isolation has been considered one of the primary causes of speciation and its genetic study has the potential to reveal the genetics of speciation. In Drosophila, the importance of courtship songs in sexual isolation between closely related species has been well investigated, but studies analysing the genetic basis of the difference in the courtship songs associated with sexual isolation are less well documented. Drosophila ananassae and Drosophila pallidosa are useful for studies of sexual isolation, because of their sympatric distribution and absence of postmating isolation. Courtship songs are known to play a crucial role in sexual isolation between these two species, and the female discrimination behaviour against the courting male has been revealed to be controlled by a very narrow region on the second chromosome. In this study we investigated the genetic basis controlling the song differences associated with their sexual isolation, using intact and wingless males with chromosomes substituted between species. The results obtained from F₁ hybrid males between these species indicate the dominance of the song characters favoured by D. pallidosa females. In addition, the results obtained from backcross F₂ males indicate that chromosome 2 had a major effect on the control of the song characters associated with sexual isolation.     

Courtship-sounds and behaviour in the four species of the Drosophila bipectinata complex

The D. bipectinata complex contains four species which are sympatric over parts of their range. They are morphologically identical and hybridize in no-choice mating situations. They have similar but distinct courtship patterns and males of all species sing two songs, long song early and short song late in courtship. Each species has a unique song profile due to differences in at least one song parameter. This is circumstantial evidence that sounds function to maintain sexual isolation within the complex.     

Morphological comparison among Drosophila lini and its two new sibling species (Diptera: Drosophilidae)

Morphological differences are investigated using several culture strains of three sibling species collected from Taiwan and Guangdong in China and Pyinoolwin and Yangon in Myanmar. Careful examination of male terminalia reveals distinguishable differences in the paramere and the aedeagal basal process among the three species. In addition, a number of quantitative characters are compared. Kruskal-Wallis tests with Bonferroni correction, which are carried out separately for each sex, detect significant differences in 15 characters, of which two are male-specific, among the three species. Canonical discriminant analysis using these characters reveals that the three species can be distinguished from each other with high confidence for both sexes. The results clearly show the presence of three good species, Drosophila (Sophophora) lini Bock & Wheeler, 1972 and its two new siblings. The new species are described as Drosophila (Sophophora) ohnishii sp. nov. from Pyinoolwin and Drosophila (Sophophora) ogumai sp. nov. from Yangon. The morphological differentiation among the three sibling species does not coincide with the degree of reproductive isolation (based on a previous study). The premating isolation pattern suggests two possibilities that premating isolation has been evolved or reinforced in sympatric populations between D. ohnishii and D. lini and between D. ohnishii and D. ogumai or that it has evolved in a very restricted local population of D. ohnishii, possibly by a few mutations.     

Latitudinal trends in wood anatomy within species and genera: case study in Cornus s.l. (Cornaceae)

Latitudinal trends in wood anatomical characters in three Asiatic species of Cornus sensu lato (s.l.) were studied and compared with those for the whole genus based on an extensive sampling covering the specific distribution ranges and the generic data from a previous study. We studied 124 specimens of C. controversa growing between 31.5° and 45.3° N, 54 of C. kousa between 24.4° and 40.5° N, and 64 of C. macrophylla between 27.8° and 41.0° N. Characters studied were vessel element length, fiber length, vessel frequency, tangential vessel diameter, and vessel grouping index. At the species level no latitudinal trends were detected throughout the distribution ranges of the species. Neither tree size, altitude, nor climatic factors had a significant correlation with wood anatomical characters. In contrast, at the genus level, latitudinal trends were significant not just for the whole genus, but for both New and Old World species groups. At the genus level, latitude and three climatic factors all had a significant correlation with wood anatomical characters, but correlation coefficients with latitude were markedly high. The difference in latitudinal trends between the genus and species levels may be due to the radiation of Cornus along paleoclimatic gradients in the early Tertiary.     

Detection of two zoonotic Babesia microti lineages, the Hobetsu and U.S. lineages, in two sympatric tick species, ixodes ovatus and Ixodes persulcatus, respectively, in Japan

The species Babesia microti, commonly found in rodents, demonstrates a high degree of genetic diversity. Three lineages, U.S., Kobe, and Hobetsu, are known to have zoonotic potential, but their tick vector(s) in Japan remains to be elucidated. We conducted a field investigation at Nemuro on Hokkaido Island and at Sumoto on Awaji Island, where up to two of the three lineages occur with similar frequencies in reservoirs. By flagging vegetation at these spots and surrounding areas, 4,010 ticks, comprising six species, were collected. A nested PCR that detects the 18S rRNA gene of Babesia species revealed that Ixodes ovatus and I. persulcatus alone were positive. Lineage-specific PCR for rRNA-positive samples demonstrated that I. ovatus and I. persulcatus carried, respectively, the Hobetsu and U.S. parasites. No Kobe-specific DNA was detected. Infected I. ovatus ticks were found at multiple sites, including Nemuro and Sumoto, with minimum infection rates (MIR) of ～12.3%. However, all I. persulcatus ticks collected within the same regions, a total of 535, were negative for the Hobetsu lineage, indicating that I. ovatus, but not I. persulcatus, was the vector for the lineage. At Nemuro, U.S. lineage was detected in 2 of 139 adult I. persulcatus ticks (MIR, 1.4%), for the first time, while 48 of I. ovatus ticks were negative for that lineage. Laboratory experiments confirmed the transmission of Hobetsu and U.S. parasites to hamsters via I. ovatus and I. persulcatus, respectively. Differences in vector capacity shown by MIRs at Nemuro, where the two species were equally likely to acquire either lineage of parasite, may explain the difference in distribution of Hobetsu throughout Japan and U.S. taxa in Nemuro. These findings are of importance in the assessment of the regional risk for babesiosis in humans.