Short-term population differences in the genetic architecture of life history traits related to sexuality in an aphid species

One of the most important factors that determine the evolutionary trajectory of a suite of traits in a population is the structure of the genetic variance-covariance matrix (G). We studied the cyclically parthenogenetic aphid Rhopalosiphum padi, whose populations exhibit two types of reproductive lineages respectively specialized in sexuality (that is, cyclically parthenogenetic lineages) and in asexuality (that is, obligate parthenogenetic lineages). We compared the quantitative genetics of life histories in these two lineage types. Our results suggest that both, the elements and the whole structure of the resulting G matrices differ in the very short term, between lineage types. This would involve the evolution toward different evolutionary optima in the same population, depending on whether sexual or asexual lineages predominate. Since sexual and asexual lineages vary seasonally in their abundance, a fluctuating selective regime has been proposed for this species, which would contribute to the maintenance of the reproductive polymorphism that these populations exhibit.     

Distribution and genetic characteristics of Ephoron shigae (Ephemeroptera: Polymitarcyidae) in Korea

Ephoron shigae is distributed in Far East Russia, Korea and Japan, where it inhabits rivers, streams and lakes. Although this mayfly is a geographically parthenogenetic species, unisex populations have been observed only in Japan. We aimed to answer the following three questions: (i) whether there is a unisex population of E. shigae in Korea; (ii) whether migration and gene flow exists between E. shigae populations in Korea and Japan; and (3) whether E. shigae populations are fragmented in Korea because of mountains running to the north and south. We observed 20 populations across Korea and investigated the sex ratio in 12 populations. Our results showed that all the populations were bisex, and we believe that there is no unisex population of E. shigae in Korea. In addition, we examined the genetic structure of E. shigae in Korea by using the COI gene. The results showed that the Korean populations were clearly divergent from the Japanese populations; the level of genetic distance between the Korean and Japanese populations was interspecific rather than intraspecific. Therefore, parthenogenetic populations of E. shigae probably belong to unisex populations that originated in Japan, while the individuals would have not migrated to Korea. The Korean populations did not show clear fragmentation by geographic barrier, although this mayfly with extremely short adult stage (up to 2 h) could be a comparatively strong disperser across Korea.     

Loss and gain of sexual reproduction in the same stick insect

The outcome of competition between different reproductive strategies within a single species can be used to infer selective advantage of the winning strategy. Where multiple populations have independently lost or gained sexual reproduction it is possible to investigate whether the advantage is contingent on local conditions. In the New Zealand stick insect Clitarchus hookeri, three populations are distinguished by recent change in reproductive strategy and we determine their likely origins. One parthenogenetic population has established in the United Kingdom and we provide evidence that sexual reproduction has been lost in this population. We identify the sexual population from which the parthenogenetic population was derived, but show that the UK females have a post‐mating barrier to fertilisation. We also demonstrate that two sexual populations have recently arisen in New Zealand within the natural range of the mtDNA lineage that otherwise characterizes parthenogenesis in this species. We infer independent origins of males at these two locations using microsatellite genotypes. In one population, a mixture of local and nonlocal alleles suggested males were the result of invasion. Males in another population were most probably the result of loss of an X chromosome that produced a male phenotype in situ. Two successful switches in reproductive strategy suggest local competitive advantage for outcrossing over parthenogenetic reproduction. Clitarchus hookeri provides remarkable evidence of repeated and rapid changes in reproductive strategy, with competitive outcomes dependent on local conditions.     

Thysanophora penicillioides includes multiple genetically diverged groups that coexist respectively in Abies mariesii forests in Japan

We investigated intraspecific diversity and genetic structures of a saprotrophic fungus--Thysanophora penicillioides--based on sequences of nuclear ribosomal internal transcribed spacer (ITS) in 15 discontinuous Abies mariesii forests of Japan. In such a well-defined morphological species, numerous unexpected ITS variations were revealed: 12 ITS sequence types detected in 254 isolates collected from 15 local populations were classified into five ITS sequence groups. Maximally, four ITS groups consisted of seven ITS types coexisting in one population. However, group 1 was dominant with approximately 65%; in particular, one haplotype, 1a, was most dominant with approximately 60% in respective populations. Therefore, few differences were recognized in genetic structure among local populations, implying that the gene flow of each lineage of the fungus occurs among local populations without geographic limitations. However, minor haplotypes in some ITS groups were found only in restricted areas, suggesting that they might expand steadily from their places of origin to neighboring A. mariesii forests. Aggregating sequence data of seven European strains and four North American strains from various substrates to those of Japanese strains, 18 ITS sequence types and 28 variable sites were recognized. They were clustered into nine lineages by phylogenetic analyses of the beta-tubulin and combined ITS and beta-tubulin datasets. According to phylogenetic species recognition by the concordance of genealogies, respective lineages correspond to phylogenetic species. Plural phylogenetic species coexist in a local population in an A. mariesii forest in Japan.     

A phylogeographical study of the Turnip mosaic virus population in East Asia reveals an 'emergent' lineage in Japan

The genetic structure of populations of Turnip mosaic virus (TuMV) in East Asia was assessed by making host range and gene sequence comparisons of 118 isolates utilizing a population genetic approach. Most, but not all, isolates collected from Brassica plants in China infected only Brassica plants, whereas those from Japan infected both Brassica and Raphanus (BR) plants. Analyses of the positions of recombination sites in five regions of the genomes (one third of the full sequence) of the many recombinant isolates were fully congruent with the results of phylogenetic analysis, and at least one recombination type pattern was shared between Chinese and Japanese populations. One lineage of nonrecombinant isolates from the basal-BR lineage was found in 2000 in Kyushu, Japan but none in China, and have since been found over the whole island. The sudden expansion of this basal-BR population was strongly supported by calculations showing the deviations from the neutral equilibrium model for the individual geographical lineages with overall lack of nucleotide diversity, and by analysis of mismatch distribution. Our study shows that the recent Chinese and Japanese TuMV isolates are part of the same population but are discrete lineages.     

Planarian mitochondria sequence heterogeneity: relationships between the type of cytochrome c oxidase subunit I gene sequence, karyotype and genital organ

Freshwater planarians Dugesia japonica from three localities were examined for cytochrome c oxidase subunit I (COI) gene sequence, karyotype and the presence of genital organ. The planarians from Mt Fujiwara in Japan were composed of two different groups; one revealed inter- and intraindividual COI gene heterogeneity, while another revealed no sequence heterogeneity. The sequence in planarians from Mt Alishan in Taiwan was homogeneous, while that from the Kenting National Park in Taiwan revealed a considerable heterogeneity. All the planarians having the homogeneous gene sequences carry the 2X karyotype and many of them had genital organs. These are assumed to belong to the sexual lineage. In contrast, almost all planarians having heterogeneous sequences carry the karyotype of either 3X plus 2X (mixoploid) or 3X, and all of them lack genital organs. These lineages are assumed to be asexual. The heterogeneity of COI gene sequences in the presumed asexual lineages would have resulted from an accumulation of mutations by repeated asexual reproduction.     

Identification of two invasive Cacopsylla chinensis (Hemiptera: Psyllidae) lineages based on two mitochondrial sequences and restriction fragment length polymorphism of cytochrome oxidase I amplicon

The occurrence of pear decline, a disease found in some pear (Pyrus spp.) orchards of Taiwan in recent years, is accompanied by an outbreak of Cacopsylla chinensis (Yang & Li). Two major morphological forms (summer and winter forms) with a variety of intermediate body color and two phylogenetic lineages of this psyllid have been described. The work herein used sequences of mitochondrial cytochrome oxidase I (COI) and 16S rDNA regions to delineate the genetic differentiation of this color-variable insect and to elucidate their relationship. Sequence divergence and phylogenetic analysis have shown that C. chinensis individuals could be divided into two lineages with 3.3 and 2.3% divergence of COI and 16S rDNA, respectively. All specimens from China were found to belong to lineage I. Restriction fragment length polymorphism analysis of COI with restriction enzymes AcuI, AseI, BccI, and FokI on 263 specimens of six populations from Taiwan produced two digestion patterns, which are in agreement with the two lineages described above. Both patterns could be found in each population, with most individuals belonging to lineage I and 5-21% of the individuals belonging to lineage II. Because these two lineages included summer as well as winter morphological forms, the lineage differentiation is apparently not related to morphological characters of this psyllid. Because the invasive records are not in favor of a sympatric differentiation, this psyllid is more likely introduced as different populations from countries in temperate regions.     

A molecular phylogeography approach to biological invasions of the New World by parthenogenetic Thiarid snails

The parthenogenetic snail Melanoides tuberculata, present in tropical fresh waters of most of the Old World before 1950, has now invaded the Neotropical area. The phylogeography of this snail was studied to evaluate the pathways and number of such invasions. Because of parthenogenetic reproduction, individuals are structured into genetical clones. Within populations from both the original and invaded areas, several morphologically distinct clones (referred to as morphs) often coexist but the amount of genetic divergence among morphs is unknown. Individuals from 27 morphs and 40 populations world-wide were sequenced at two mitochondrial genes (12S and 16S). Our phylogenetic reconstruction suggests that (i) most of the morphological variation observed in the New World predates invasion, (ii) at least six independent introductions have occurred, and (iii) invasive clones are found throughout most of the phylogenetic tree and do not come from a particular region of the area of origin. Two ideas are discussed in the light of these results. The first lies with the specificities of parthenogenesis in an invasion context. While in sexual species, independently introduced populations eventually merge into a single invasive population, in a parthenogenetic species independently introduced clones have distinct invasion dynamics and possibly exclude each other. Second, although repeated invasions in Melanoides may have an impact on indigenous molluscan faunas, their most likely effect is the world-wide homogenization of the invasive taxon itself.     

Morphological stasis and phylogenetic relationships in Tadpole shrimps, Triops (Crustacea: Notostraca)

The tadpole shrimp (Triops) is a well-known 'living fossil' whose fundamental morphology has been unchanged for over 170 million years. Thus, tadpole shrimps are suitable subjects for the study of morphological stasis. We were able to obtain samples of three species of Triops (T. granarius, T longicaudatus and T. cancriformis) from four regions in Japan. Taxonomic species were identified by diagnostic morphology. We inferred phylogenetic relationships between individual samples using mitochondrial 16S rRNA. Carapace shapes were compared among populations using shape coordinate methods. The phylogeny inferred from mtDNA shows that T. granarius is phylogenetically more similar to T. longicaudatus than to T. cancriformis. mtDNA sequences did not differ among the populations of T granarius. However, there were two distinct phylogenetic species within T. longicaudatus. In spite of the similarity in fundamental morphological characteristics among Triops species, mtDNA sequences of Triops showed marked differences among the four phylogenetic species. Among the populations of T. granarius , the carapace shape of the Fukuoka population was significantly different from those of other populations of T. granarius. The carapace shape of the Kagawa population of T. longicaudatus was more similar to those of the Shizuoka and Kagawa populations of T. granarius. The shape of the carapace of T. cancriformis was significantly different from those of T. granarius and T longicaudatus. Thus, taxonomic species, phylogenetic species and populations with similar carapace morphology did not correspond with each other. The present results indicate that most of the morphological change did not occur at the time of speciation (lineage separation) and that morphological stases are important evolutionary patterns, but they are not species-level properties.     

Parthenogenetic populations of the freshwater snail Campeloma limum occupy habitats with fewer environmental stressors than their sexual counterparts

1. Sexual organisms should have half the reproductive rate of their parthenogenetic counterparts (i.e. twofold cost of sex), so the plethora of sexual species relative to parthenogenetic species remains an evolutionary paradox. The rarity of parthenogenesis may in part be due to the accumulation of deleterious mutations. Indeed, parthenogenetic populations of the freshwater snail Campeloma limum have a greater mutation load relative to sexual populations of C. limum, although this does not directly affect their reproductive fitness. We hypothesise that although parthenogenesis has no direct effect on fitness in C. limum, mutation accumulation and environmental stress act synergistically to limit the distribution of parthenogenetic populations. 2. We evaluated this hypothesis, predicting that parthenogenetic populations of C. limum would inhabit sites with fewer environmental stressors than their sexual counterparts. We collected water quality, population density and individual size data at multiple time points from eight parthenogenetic and five sexual populations in the south‐eastern United States (Georgia and South Carolina). 3. Consistent with our hypothesis, sexual populations of C. limum inhabited poorer‐quality areas (sites with significantly lower dissolved oxygen and significantly more faecal coliform bacteria) than parthenogenetic populations. Despite these stressors, sexual populations still exhibited significantly higher population density than parthenogenetic populations. 4. Our findings support the hypothesis that mutation‐laden parthenogenetic C. limum populations occupy habitats with fewer environmental stressors relative to their sexual counterparts. Moreover, sexual C. limum populations inhabit lower‐quality habitats where they can presumably evade the twofold cost of sex in the absence of competition from their parthenogenetic counterparts.     

Russula velenovskyi new to Japan,with phylogenetic implications of Russula species between Japanese subalpine forests and Northern Europe

To compare morphological characters and phylogenetic placement between Japanese and European Russula, 32 specimens of 12 species were collected from Japanese subalpine forests and Northern Europe. Several sequences of nrDNA ITS region (ITS) of these Russula species were obtained. High homological similarities were shown between ITS sequences of several Russula samples collected from Japanese subalpine forests, Europe and North America. These facts show distribution of the same Russula species among these areas. From morphological observations and phylogenetic analyses, two same Russula species, R. velenovskyi, and R. decolorans are found in Japan, Europe and North America. Of these, R. velenovskyi collected from Mt. Fuji, Mt. Nyukasa and Mt. Tateshina in mountainous area of central Honshu is reported as a new Japanese record.     

Salinity effects on maturation, reproductive and life span characteristics of four Egyptian Artemia populations (International Study on Artemia. LXVIII)

Three parthenogenetic Artemia populations, i.e. two coastal from Borg El-Arab and El-Max saltworks and one from the inland Qarun Lake, and a bisexual strain (Artemia salina) from the inland carbonate lake of Wadi El-Natrun, all from upper Egypt were assayed for their response in 5 salinities (i.e. 35, 80, 120, 150 and 200 g l^?1). The experimental procedure was carried out under laboratory conditions, where the effects of salinity on maturation and ten reproductive and life span characteristics were investigated. The parthenogenetic Egyptian populations are more euryhaline compared to the bisexual one. The two coastal parthenogenetic populations appeared to be very similar in maturation rate and reproductive output at all salinities tested. The inland asexual strain showed a different reproductive response to the elevation of salinity from the two coastal populations. Discriminant function analysis has proven to be a useful tool in determining the differential response of closely related Artemia populations. The bisexual population showed significantly lower reproductive output compared to the parthenogenetic ones and performed best at 35 g l^?1; this is the first record of an A. salina population inhabiting a carbonate lake. These findings may provide valuable information on Artemia biodiversity in an area where very little is known. %     

Parthenogenetic female populations in the brown alga Scytosiphon lomentaria (Scytosiphonaceae,Ectocarpales): decay of a sexual trait and acquisition of asexual traits

In isogamous brown algae, the sexuality of populations needs to be tested by laboratory crossing experiments, as the sexes of gametophytes are morphologically indistinguishable. In some cases, gamete fusion is not observed and the precise reproductive mode of the populations is unknown. In the isogamous brown alga Scytosiphon lomentaria in Japan, both asexual (gamete fusion is unobservable) and sexual populations (gamete fusion is observable) have been reported. In order to elucidate the reproductive mode of asexual populations in this species, we used PCR‐based sex markers to investigate the sex ratio of three asexual and two sexual field populations. The markers indicated that the asexual populations consisted only of female individuals, whereas sexual populations are composed of both males and females. In culture, female gametes of most strains from asexual populations were able to fuse with male gametes; however, they had little to no detectable sexual pheromones, significantly larger cell sizes, and more rapid parthenogenetic development compared to female/male gametes from sexual populations. Investigations of sporophytic stages in the field indicated that alternation of gametophytic and parthenosporophytic stages occur in an asexual population. These results indicate that the S. lomentaria asexual populations are female populations that lack sexual reproduction and reproduce parthenogenetically. It is likely that females in the asexual populations have reduced a sexual trait (pheromone production) and have acquired asexual traits (larger gamete sizes and rapid parthenogenetic development).     

Reproductive modes,ploidy distribution,and supernumerary chromosome frequencies of the flatworm Polycelis nigra (Platyhelminthes: Tricladida)

The hermaphroditic flatworm, Polycelis nigra, is characterized by two reproductive biotypes which differ with respect to ploidy; sexual individuals are diploid (n = 8, 2× = 16) and pseudogamous parthenogenetic individuals are polyploid (typically 3×). We have collected and karyotyped individuals from 15 sampling sites (13 in mid to northern Italy, one in Great Britain and one in The Netherlands). We found that biotypes can exist alone or in sympatry, and identified purely diploid, mixed diploid-polyploid, and purely polyploid populations. Karyotype data show that in addition to the normal autosome complement, B chromosomes of differing morphology as well as stable aneuploid chromosomes (extra-A) were found almost exclusively in polyploids (11 of 12 sites). We extensively sampled Lago di Toblino (northern Italy), a pure polyploid population characterized by a submetacentric to metacentric, mitotically stable B chromosome, as well as a stable extra-A chromosome. Here, individuals having 1–3 B chromosomes were more abundant (61%) than those having no B's, implying that B chromosome infection has little detrimental effect when occurring in low numbers. Furthermore, 66% of individuals from this population possessed extra-A chromosomes, although it is unclear whether these elements are aneuploid autosomes or B chromosomes of different morphology. The ubiquity of these chromosomes, within asexuals in particular, is suggestive of a correlation between the origination of the elements and the evolution of polyploidy, or may reflect increased tolerance of parthenogenetic genomes to aneuploidy. This revised version was published online in July 2006 with corrections to the Cover Date.     

Using multilocus approach to uncover cryptic diversity within Pseudotrapelus lizards from Saudi Arabia

The agamid Pseudotrapelus lizards inhabit the mountainous areas of the Arabian Peninsula and eastern North Africa. Currently six Pseudotrapelus species are recognised, though diagnostic morphological characters are still lacking, creating great difficulty in describing new species. Recently, two specimens of Pseudotrapelus were collected from the vicinity of Riyadh in central Saudi Arabia, an area that was not sampled in previous phylogenetic studies. In here we used both mitochondrial and nuclear data to investigate the phylogenetic position of the new samples, and assess their phylogenetic relationships with the other recognised species of Pseudotrapelus from across the distribution range of the genus. We used a multilocus approach of haplotype networks, concatenated datasets and species trees, performed mitochondrial and nuclear species delimitation analyses, and estimated divergence times. In general, our results support previous molecular studies and uncover the presence of cryptic diversity within Pseudotrapelus. The phylogenetic structure of the genus is of two major clades and within them seven distinct, delimited phylogenetic groups belonging to the six recognised species and the seventh to the individuals from Riyadh. The Riyadh specimens were distinct in all analyses performed. We suggest that the new specimens from the Riyadh area are a distinct lineage, forming a clade with their phylogenetic relatives, P. sinaitus and P. chlodnickii. The clade formed by these three species diverged during the Late Miocene around 6.4 Ma, with cladogenesis possibly facilitated by vicariance and isolation caused due to climatic fluctuations and the progression of sandy areas. Our results suggest further morphological research is necessary to revise the taxonomic status of this lineage and of the entire genus.     

The post-natal development of holocrine epidermal specializations in gekkonid lizards.

The abdominal escutcheon, and certain aspects of pre-anal organ morphology, have been studied in Sphaerodactylus spp. and Gekko vittatus respectively. These epidermal modifications are male characteristics. The sphaerodactyline escutcheon becomes larger by the peripheral addition of specialized scales with increasing size of the individuals: this relationship is much more clearcut in S. cinereus than in the notatus species group (sensu Shreves, '68), and the possible reasons for this are discussed. The number of pre-anal organs varies between populations of G. vittatus, but within populations remains constant throughout life. Individual organs increase steadily in size throughout life. These data are discussed with reference to current interpretations of gekkonid gland evolution, and of factors controlling epidermal cell proliferation and differentiation.     

Cheater genotypes in the parthenogenetic ant Pristomyrmex punctatus

Cooperation is subject to cheating strategies that exploit the benefits of cooperation without paying the fair costs, and it has been a major goal of evolutionary biology to explain the origin and maintenance of cooperation against such cheaters. Here, we report that cheater genotypes indeed coexist in field colonies of a social insect, the parthenogenetic ant Pristomyrmex punctatus. The life history of this species is exceptional, in that there is no reproductive division of labour: all females fulfil both reproduction and cooperative tasks. Previous studies reported sporadic occurrence of larger individuals when compared with their nest-mates. These larger ants lay more eggs and hardly take part in cooperative tasks, resulting in lower fitness of the whole colony. Population genetic analysis showed that at least some of these large-bodied individuals form a genetically distinct lineage, isolated from cooperators by parthenogenesis. A phylogenetic study confirmed that this cheater lineage originated intraspecifically. Coexistence of cheaters and cooperators in this species provides a good model system to investigate the evolution of cooperation in nature.     

Trophic ecology of East African caecilians (Amphibia: Gymnophiona), and their impact on forest soil invertebrates

Diets of the syntopic caecilian amphibians Boulengerula boulengeri and Scolecomorphus vittatus were studied from gut content analyses of 108 specimens collected from Nilo Forest Reserve, East Usambara Mountains, Tanzania. Head length and width relative to body length is greater in male B. boulengeri than in females, but no such dimorphism is apparent in S. vittatus . No differences, other than females consuming disproportionately more ants and invertebrate eggs and fewer termites than males, occur in the diets of B. boulengeri . The gut contents of S. vittatus are dominated by large, surface-active earthworms, in contrast to those of B. boulengeri which contained smaller, endogeic earthworms and a much greater number and diversity of soil arthropods (particularly termites, ants and Diptera larvae). These dietary characteristics underline differences in microhabitat use between these caecilian species and thus corroborate a previously proposed hypothesis of niche separation. The ecological impact of caecilians and other limbless endogeic vertebrates is little studied and poorly known. Approximations based on current knowledge are not clear, but are interpreted as indicating that predation by caecilians is unlikely to be among the most important factors influencing population densities of soil-dwelling termites and perhaps also ants: two groups recognized as soil ecosystem engineers.     

Modelling the lineage of a growing population as an age-dependent branching process

A new model for population growth as a branching process is described. It requires knowledge of the population's age-specific survivorship and fecundity curves. The total population size at any time is calculated by summing across all branches of the tree representing the population's lineage and thus the model also allows the lineage of the developing population to be traced. Use of the model is demonstrated by a hypothetical example. The model could be of particular value in tracing the phylogenetic development of populations and/or species.     

Mitochondrial DNA Diversity of Monopterus albus from the Sichuan Basin of China

Genetic diversity of Monopterus albus in a wild population in the Sichuan Basin was investigated by employing mtDNA polymorphisms. The mitochondrial control region was sequenced for a total of 59 individuals collected from four localities. These sequences (553 bp) belonged to 13 haplotypes defined by eight polymorphic sites. The haplotypes were clustered into one lineage, suggesting that Monopterus albus from the Sichuan Basin might originate from a single ancestor in maternal lineages. The genetic differentiation between the population from Zhongxian in Chongqing city and the other three populations, from Mianyang, Longchang, and Ya'an in Sichuan province (F (ST): 0.29167-0.55126), was higher than that among the latter three (F (ST): -0.04196 to 0.05984). Little genetic differentiation was found in the three populations from Sichuan province (Nm: -6.21 to 3.93), and the Zhongxian population also showed lower genetic heterogeneity (P (i): 0.00077 +/- 0.00027; H (d): 0.404 +/- 0.130). The combination of limited variation and weak phylogenetic structure revealed that the Monopterus albus from the Sichuan Basin exhibited lower mtDNA diversity and should probably be treated as a monogroup genetically.