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1.
The cereal rust mite, Abacarus hystrix, is one of the most notable among mites causing losses in cultivated grasslands. It is one of a few eriophyoid species for which a broad host range has been reported. Recent studies, however, have shown that host specialization is very likely in this species. For two populations of A. hystrix (one inhabiting perennial ryegrass, the second inhabiting quackgrass), host-associated differences correlated with strong host fidelity, distinct phenotypes and reproductive barriers have been found. In the present study, the ability of wheat colonization by quackgrass- and ryegrass-adapted cereal rust mite was studied. The hypothesis that the potential for wheat colonization by the quackgrass strain is more likely was tested by comparing the colonization performance (assessed by female survival and fecundity) of quackgrass- and ryegrass-associated A. hystrix on their familiar hosts and on wheat. The ryegrass population had no success in wheat colonization (expressed by extremely low fecundity and female survival). Fecundity and survival of quackgrass strain were similar on wheat and the familiar host, or even higher on wheat. Phylogenetic similarity of quackgrass and wheat is discussed as a possible factor that might influence such patterns of host colonization. Since A. hystrix is the only vector of the ryegrass mosaic virus (RgMV), the presented results may be helpful in explaining the inability of RgMV to successfully infest wheat. The conclusions are that (i) quackgrass- and ryegrass-adapted strains of the cereal rust mite have different physiological host ranges and (ii) phylogenetic relationships between host plant species appear to be drivers for host specialization in this mite species.  相似文献   

2.
For phytophagous arthropods, host acceptance behavior is a key character responsible for host plant specialization. The grain rust mite, Abacarus hystrix (Nalepa), is an obligately phytophagous, polyphagous eriophyid mite recorded from at least 70 grass species. In this study, the hypothesis that two host populations of this mite (one collected from quackgrass and the other from ryegrass) are highly host-specific was tested using behavioral data. For this purpose, female behavior when exposed to familiar and novel host plants was observed in no-choice cross experiments. Altogether, 13 variables were used to describe mite behavior. Data were subjected to principal component analysis, and host acceptance behavior was subsequently tested with generalized estimating equations (GEE). Distinct variation in female behavior between familiar and novel hosts was observed. Females from neither population accepted novel hosts. This was recorded as significant differences in the occupation of and overall activity on particular plant parts. On their familiar host, females were not active and showed little tendency to move. On novel hosts females were more active and mobile, spending more time walking, running, and climbing on the whole plant surface and showing a tendency to disperse. Other differences in behavior between studied populations were also observed. Thus, the results suggest that mites of these two studied populations (1) differ in their behaviors during plant exploitation and (2) can quickly distinguish between their familiar host and an unfamiliar host used by a conspecific. These findings support the hypothesis of narrow host specialization of ryegrass and quackgrass populations of this highly polyphagous species.  相似文献   

3.
When species hybridize, one F1 hybrid cross type often predominates. Such asymmetry can arise from differences in a variety of reproductive barriers, but the relative roles and concordance of pre-mating, post-mating prezygotic, and post-zygotic barriers in producing these biases in natural animal populations have not been widely investigated. Here, we study a population of predominantly F1 hybrids between two killifish species (Fundulus heteroclitus and F. diaphanus) in which >95% of F1 hybrids have F. diaphanus mothers and F. heteroclitus fathers (D♀ × H♂). To determine why F. heteroclitus × F. diaphanus F1 hybrids (H♀ × D♂) are so rare, we tested for asymmetry in pre-mating reproductive barriers (female preference and male aggression) at a common salinity (10 ppt) and post-mating, pre-zygotic (fertilization success) and post-zygotic (embryonic development time and hatching success) reproductive barriers at a range of ecologically relevant salinities (0, 5, 10, and 15 ppt). We found that F. heteroclitus females preferred conspecific males, whereas F. diaphanus females did not, matching the observed cross bias in the wild. Naturally rare H♀ × D♂ crosses also had lower fertilization success than all other cross types, and a lower hatching success than the prevalent D♀ × H♂ crosses at the salinity found in the hybrid zone centre (10 ppt). Furthermore, the naturally predominant D♀ × H♂ crosses had a higher hatching success than F. diaphanus crosses at 10 ppt, which may further increase their relative abundance. The present study suggests that a combination of incomplete mating, post-mating pre-zygotic and post-zygotic reproductive isolating mechanisms act in concert to produce hybrid asymmetry in this system.  相似文献   

4.
Reinforcement is the process by which selection favors traits that decrease mating between two incipient species in response to costly mating or the production of maladapted hybrids, causing the evolution of greater reproductive isolation between emerging species. I have studied a pair of orchids, Neotinea tridentata and N. ustulata, to examine the level of postmating pre- and post-zygotic isolating mechanisms that maintain these species, and the degree to which the boundary may still be permeable to gene flow. In this study, I performed pollen tube growth rate experiments and I investigated pre- and post-zygotic barriers by performing hand pollination experiments in order to evaluate fruit set, embryonate seed set and seed germination rates by intra- and interspecific crosses. Fruit set, the percentage of embryonate seeds and germinability of interspecific crosses were reduced compared to intraspecific pollinations, showing significant differences between sympatric and allopatric populations. While in allopatric populations the post-pollination isolation index ranged between 0.40 and 0.11, in sympatric populations orchid pairs showed total isolation due to post-pollination prezygotic barriers, guaranteed at the level of pollen–stigma interactions. Indeed, in sympatric populations, pollen tubes reached the ovary after 24 h in only 8 out of 45 plants; in the remaining cases, the pollen tubes did not enter the ovary, and thus no fruit set occurred. This pair of orchids is characterized by postmating pre-zygotic reproductive isolation in sympatric populations that prevents the formation of hybrids. This mechanism of speciation, starting in allopatry and triggering the reinforcement mechanisms of reproductive isolation in secondary sympatry, is the most likely explanation for the pattern of evolutionary transitions found in this pair of orchids.  相似文献   

5.
The objective of the present research is to study the reproductive compatibility between populations of predatory mites previously identified as Euseius concordis (Chant) based on morphological characteristics. Colonies of these mite populations were established in the lab with specimens collected from different localities and host plants. Reproductive compatibility was evaluated through crosses and backcrosses within and between populations and the subsequent observation of females' oviposition, over a period of 10 days. The levels of oviposition obtained in the crosses between individuals from the same population were higher than those obtained in the crosses between individuals from different populations. Results indicate the occurrence of post-mating reproductive incompatibility between the mite population from Petrolina and the other populations studied. Crosses and backcrosses between populations involving female mites from Petrolina did not produce offspring, although endospermatophores were present inside the spermathecas of those females. Oviposition was reduced, and only sons were obtained, in crosses between populations with males from Petrolina. Crosses of females from Pontes e Lacerda and males from Jaguariúna and vice versa produced only male progeny. Our results established that the populations originating from Arroio do Meio, Pontes e Lacerda, Jaguariúna and Viçosa, are reproductively compatible. However, the latter populations and the population from Petrolina are genetically isolated. Based on these results we suggest that more cytological and genetic studies are needed to establish if this reproductive isolation represents a species barrier.  相似文献   

6.
The majority of eriophyoid mites are highly host specific and restricted to a narrow range of acceptable host plant species. The cereal rust mite, Abacarus hystrix was considered to be one of a few exceptions among them and has been found to be using a relatively wide host range. Since this species is a vagrant, inhabiting short-lived plants and aerially dispersing, it has commonly been considered to be a host generalist. Here the opposite hypothesis is tested, that host populations of A. hystrix are specialized on their local host plants and may represent host races. For this purpose, females from two host populations (quack grass, Agropyron repens and ryegrass, Lolium perenne) were transferred, and subsequently reared, on their normal (grass species from which females came from) and novel (other grass species) hosts. The female's fitness was assessed by survival and fecundity on the normal and novel host. Females of both populations had no success in the colonization of the novel host. They survived significantly better and had significantly higher fecundity on their normal host than on the novel one. These findings correspond with observations on host-dependent phenotype variability and host acceptance. The presence of locally specialized host populations in A. hystrix may be evidence for high host specificity among eriophyoids and the viruses they transmit. The main conclusion is that A. hystrix, which so far has been considered as a host generalist, in fact may be a complex species consisting of highly specialized host races.  相似文献   

7.
Developmental success on 47 species of plant and reproductive compatibility were studied for four strains of the European red mite,Panonychus ulmi (Koch), collected from dwarf bamboo, elm and apple trees. The host range was variable in those strains and there was no plant species on which all four strains were able to reach maturity.Intra-populational crosses in the dwarf-bamboo strain and the two apple strains of mites, as well as crosses between the two apple strains, gave both female and male progeny. Inter-populational crosses between the dwarf-bamboo and apple strains produced only male progeny, as in the case where females of each strain were kept virgin, indicating that these two were reproductively incompatible and had an arrhenotokous reproductive system. When females of the elm strain were crossed with males of the elm, dwarf-bamboo or apple strain, no males were produced, as in the case for virgin females of the elm strain which had a thelytokous reproductive system. It is thus concluded that the strains derived from different host species were reproductively incompatible with one another.  相似文献   

8.
Theory suggests that post-zygotic reproductive isolation of polyploids from their diploid progenitors is often caused by developmental disorder in the endosperm of hybrid seeds. Yet, this so-called triploid block is increasingly recognized to be less strong than initially assumed, indicating that other isolation mechanisms are needed to explain reproductive isolation of polyploids and diploids. In the present study, the extent of inter-ploidy block was quantified based on crossing experiments between closely related diploid, tetraploid, and hexaploid species from the genus Leucanthemum Mill. (Compositae, Anthemideae). Seed set and viability of seeds obtained from inter-cytotype crosses were measured and compared to fertilities of intra-cytotype crosses. Although inter-ploidy block was observed when diploids acted as pollen donors, the main observation was that all inter-ploidy crosses were capable of producing viable offspring. By contrast, flow cytometrical analysis of 233 individuals from natural populations did not reveal the presence of any unambiguous cross-ploidy offspring in the field. Hence, the results of the present study demonstrate that inter-cytotype mating may be rare even though inter-ploidy block is weak. Consequently, it can be assumed that pre-zygotic barriers and reduced fitness of inter-cytotype hybrids play a decisive role in the reproductive isolation of polyploid Leucanthemum species.  相似文献   

9.
Mediterranean orchids that grow in admixed, co-flowering populations, and frequently show hybrid progenies are interesting to use to study the nature and the strength of post-zygotic barriers. However, examination of pre- and post-pollination pre-zygotic isolating mechanisms requires sympatric, co-flowering species pairs that do not produce hybrid swarms. In this study, we analyzed a contact zone between Orchis italica and O. papilionacea, in which hybrid forms have never been reported, although hybridization between members of their groups of appurtenance has been signaled. We investigated pre-pollination barriers observing the floral phenology of both species and identified pollinators by means of molecular analysis of pollinaria collected on the insects captured in the study site. Post-pollination barriers were tested performing manual crosses in order to evaluate pollen germination/pollen tube growth in vivo and fruit and seed formation. Floral phenologies of O. italica and O. papilionacea display nearly overlapping trends, and two common pollinators have been identified by molecular analysis of pollinaria. Thus, pre-pollination barriers are very weak or nonexistent. Bidirectional crosses have shown that the growth of heterospecific pollen tubes is fully blocked in stigmatic cell layers. Since no fruit formation was detected in bidirectional interspecific crosses, we assume that reproductive isolation between the examined species is fully guaranteed by post-pollination pre-zygotic mechanisms acting at stigmatic level. Such condition has been rarely described and may mask the potential action of post-zygotic mechanisms.  相似文献   

10.
Wolbachia are cytoplasmically transmitted bacteria that infect several species of mites. In the two-spotted spider mite Tetranychus urticae Koch this symbiont can induce reproductive incompatibility. Wolbachia-induced reproductive incompatibility is observed in crosses between Wolbachia-infected (W) males and uninfected (U) females. This incompatibility is expressed in F1 broods as male-biased sex ratios, an effect called cytoplasmic incompatibility (CI). However, in the two-spotted spider mite, Wolbachia-induced reproductive incompatibility may extend to the F2: broods of virgin F1 females from U×W crosses sometimes suffer increased mortality rates. This F2 effect is called hybrid breakdown (HB). Several isofemale lines derived from mites collected from rose and cucumber plants had been previously tested for CI. Here we report on the results obtained for HB.  相似文献   

11.
Cardinium, a bacterium from the Bacteroidetes group, is associated with reproductive manipulations such as cytoplasmic incompatibility, parthenogenesis, and feminization in some arthropod species. We, and others, have shown that Cardinium, but not Wolbachia, is an endosymbiont in some populations of Metaseiulus occidentalis, a phytoseiid mite that is an important predator of spider mite pests of agricultural crops. However, the precise biological effects that Cardinium may have on M. occidentalis remain unclear. In this study we show, in two sets of crosses between different colonies of Cardinium-free (C-) M. occidentalis females and Cardinium-containing (C+) males, that fecundity was reduced in parental females, F1 progeny survival rates were reduced, and fewer female progeny were produced when compared to the reciprocal and control crosses. There were no differences in these attributes in the reciprocal and two control crosses. Cardinium was transmitted maternally but there was no observed paternal transmission. Finally, Cardinium did not cause asexual (thelytoky) reproduction in M. occidentalis. Thus, Cardinium is associated with nonreciprocal reproductive incompatibility in M. occidentalis and our results support the hypothesis that Cardinium is a reproductive parasite in this agriculturally important predator. Cardinium may therefore affect the evolution and ecology of M. occidentalis and biological control efforts using this mite.  相似文献   

12.
The most common post-zygotic isolation mechanism between populations of the phytophagous mite Tetranychus urticae is 'hybrid breakdown', i.e. when individuals from two different populations are crossed, F1 hybrid females are produced, but F2 recombinant male offspring suffer increased mortality. Two-spotted spider mites collected from two populations, one on rose and the other on cucumber plants, were infected with Wolbachia bacteria. These bacteria may induce cytoplasmic incompatibility in their hosts: uninfected (U) females become reproductively incompatible with infected (W) males. We report on the effect of Wolbachia infections in intra- and interstrain crosses on (i) F1 mortality and sex ratios (a test for cytoplasmic incompatibility), and (ii) the number of haploid offspring and mortality in clutches of F1 virgins (a test for hybrid breakdown). U x W crosses within the rose strain exhibited partial cvtoplasmic incompatibility. More interestingly, F2 males suffered increased mortality, a result identical to the hybrid breakdown phenomenon. The experiments were repeated using females from the cucumber strain. In interstrain U x W and U x U crosses, hybrid breakdown was much stronger in the former (80 versus 26%). This is the first report of a Wolbachia infection causing a hybrid breakdown phenotype. Our results show that Wolbhachia infections can contribute to reproductive incompatibility between populations of T. urticae.  相似文献   

13.
Haemonchus contortus, in common with many nematode species, has extremely high levels of genetic variation within and between field populations derived from distant geographical locations. MHco10(CAVR), MHco3(ISE) and MHco4(WRS) are genetically divergent H. contortus strains, originally derived from Australia, Kenya and South Africa, respectively, that have been maintained by numerous rounds of in vivo experimental infection of sheep. In order to explore potential pre-zygotic competition or post-zygotic incompatibility between the strains, we have investigated the ability of MHco10(CAVR) to interbreed with either MHco3(ISE) or MHco4(WRS) during dual strain co-infections. Sheep were experimentally co-infected with 4000 infective larvae (L3) per os of the MHco10(CAVR) strain and an equal number of either the MHco3(ISE) or the MHco4(WRS) strain L3. The adult worm establishement rates and the proportions of F1 progeny resulting from intra- and inter-strain mating events were determined by admixture analysis of microsatellite multi-locus genotypes. Although there was no difference in adult worm establishment rates, the proportions of F1 progeny of both the MHco10(CAVR) × MHco3(ISE) and MHco10(CAVR) × MHco4(WRS) dual strain co-infections departed from Mendelian expectations. The proportions of inter-strain hybrid F1 progeny were lower than the expected 50%, suggesting either pre-zygotic competition or post-zygotic incompatibility between the co-infecting strains. To investigate this further, both eggs and hatched L1 of broods from single adult female worms recovered from each dual co-infection were genotyped. Unhatched eggs from the broods revealed no inter-strain hybrid genotype deficit, suggesting there is no pre-zygotic competition between the strains. In contrast, there was a deficit in L1 inter-strain hybrid genotypes in the broods derived from MHco3(ISE) or MHco4(WRS) maternal parents, but not from MHco10(CAVR) maternal parents. This suggests that hybrid progeny of MHco10(CAVR) paternal parents have reduced post-zygotic development and/or viability consistent with incipient speciation of the MHco10(CAVR) strain. The presence of mating barriers between allopatric H. contortus strains has important implications for parasite ecology, including the ability of newly introduced anthelmintic-resistant parasite populations to compete and interbreed with populations already established in a region.  相似文献   

14.
Sequence variation of the complete second internal transcribed spacer (ITS2, 445 bp) of nuclear ribosomal DNA and part of the mitochondrial cytochrome oxidase I gene (COI, 350 bp) was examined in Amphitetranychus viennensis (Zacher) mites (Acari:Tetranychidae) from four French and four Japanese locations. Sequence analysis consistently revealed the separation of the samples in two major groups: French mites differed from Japanese by 3.8–4.1% of the nucleotide divergence in COI sequences. These two groups also displayed distinct ITS2 consensus sequences (2.1% nucleotide divergence). A few variations, not affecting the diagnostic sites around the consensus sequence, were revealed among cloned copies of the same individual. Reciprocal crosses and backcrosses between one French and two Japanese populations disclosed strong reproductive incompatibility. However, fertile hybrid females were obtained, indicating the conspecificity of the tested mites. Despite the presence of Wolbachia in the French strain, but not in the Japanese ones, our crosses did not display the unidirectional incompatibility typically produced by this microorganism, but rather a bidirectional – although asymmetrical – incompatibility pattern. The post-zygotic incompatibilities in A. viennensis cannot be explained by the presence of Wolbachia but to some extent by mite genome divergence resulting from limited gene exchange between allopatric populations. Experiments of Wolbachia elimination by antibiotic treatment and subsequent crosses with cured strains are still needed to fully understand the reproductive incompatibility patterns in this mite species.  相似文献   

15.
We assessed prezygotic (probability of spawning) and postzygotic (hatching success) reproductive isolation among the three ecologically and morphologically similar species in the Fundulus notatus species complex. We employed a multi-generation breeding experiment to test the hypotheses that karyotypic differences, body size differences, or geographic isolation among populations will increase pre or postzygotic reproductive barriers. Overall, prezygotic barriers were strong and postzygotic barriers weak in crosses of non-hybrid heterospecifics (F1 hybrid crosses) while prezygotic barriers were weaker and postzygotic barriers stronger in crosses involving hybrid individuals (F2 hybrid crosses and backcrosses). Prezygotic barriers among the two smaller species (Fundulus notatus and F. euryzonus) broke down rapidly; first generation hybrids spawned (F2 hybrid crosses and backcrosses) as frequently as parental forms in intraspecific crosses. There was no increase in postzygotic barriers among species with cytogenetic differences. There were increased prezygotic, but not postzygotic, barriers among geographically isolated populations of one species. While pure males and females were just as likely to spawn with hybrids, some types of hybrid females suffered from increased sterility, but not inviability, over hybrid males. Female sterility was only seen in hybrids with a Fundulus euryzonus parent, while other female hybrids produced viable eggs.  相似文献   

16.
Data on patterns of variation within hybrid zones, combined with studies of life history, mate choice, and hybrid performance, allow estimates of the contribution of different pre-zygotic and post-zygotic barriers to reproductive isolation. We examine the role of behavioural barriers to gene exchange in the maintenance of a hybrid zone between North American field crickets Gryllus firmus and Gryllus pennsylvanicus . We consider these barriers in the context of previous studies that documented temporal and ecological isolation and a one-way post-mating incompatibility (i.e. G. firmus females do not produce offspring when they mate only with heterospecific males). Based on no-choice mating experiments in the laboratory, we demonstrate strong behavioural pre-mating barriers between the two species, but no apparent fecundity or fertility costs for G. firmus females when they mate with both conspecific and heterospecific males. Furthermore, we show that G. firmus females do not discriminate between hybrids and conspecifics, whereas G. pennsylvanicus females do. This observation could explain the asymmetric allele introgression observed in the hybrid zone. We also document a failure of heterospecific males to induce normal oviposition in G. firmus females, which may be due to rapid evolution of accessory gland proteins and may serve as an additional barrier to gene exchange.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 390–402.  相似文献   

17.

Background and Aims

The maintenance of species boundaries in sympatric populations of closely related species requires some kind of reproductive isolation that limits gene flow among species and/or prevents the production of viable progeny. Because in orchids mycorrhizal fungi are needed for seed germination and subsequent seedling establishment, orchid–mycorrhizal associations may be involved in acting as a post-mating barrier.

Methods

We investigated the strength of post-mating barriers up to the seed germination stage acting between three closely related Orchis species (Orchis anthropophora, O. militaris and O. purpurea) and studied the role of mycorrhizal fungi in hybridization by burying seed packets of pure and hybrid seeds. After retrieval and assessment of seed germination, the fungi associating with protocorms originating from hybrid and pure seeds were determined and compared with those associating with adult individuals using DNA array technology.

Results

Whereas pre-zygotic post-mating barriers were rather weak in most crosses, post-zygotic post-mating barriers were stronger, particularly when O. purpurea was crossed with O. anthropophora. Germination trials in the field showed that seed germination percentages of hybrid seeds were in most cases lower than those originating from pure crosses. In all species pair combinations, total post-mating reproductive isolation was asymmetric. Protocorms associated with a smaller range of fungal symbionts than adult plants, but there was considerable overlap in mycorrhizal associations between protocorms and their respective parents.

Conclusions

Our results suggest that mycorrhizal associations contribute little to reproductive isolation. Pre-mating barriers are probably the main factors determining hybridization rates between the investigated species.  相似文献   

18.
Selection along environmental gradients can drive reproductive isolation and speciation. Among fishes, salinity is a major factor limiting species distributions, and despite its importance in generating species diversity, speciation events between marine and freshwater are rare. Here, we tested for mechanisms of reproductive isolation between locally adapted freshwater and brackish water-native populations of killifish, Fundulus heteroclitus, from either side of a hybrid zone along a salinity gradient. There was evidence for pre-zygotic endogenous reproductive isolation with reduced fertilization success between crosses of freshwater-native males and brackish water-native females. Exogenous pre-zygotic isolation was also present where females had highest fertilization in their native salinity. We used a replicated mass spawning design to test for mate choice in both brackish and fresh water. After genotyping 187 parents and 2523 offspring at 2347 SNPs across the genome, 85% of offspring were successfully assign to their parents. However, no reinforcing mate choice was observed. These results therefore demonstrate emerging, yet limited, reproductive isolation and incipient speciation across a marine to freshwater salinity gradient and suggest that both endogenous and exogenous mechanisms, but not assortative mating, contribute to divergence.  相似文献   

19.
Abstract: Quantitative characteristics of the occurrence of eriophyoid mites on wheat and associated grasses were studied. Three groups of hosts: cultivated wheat ( Triticum aestivum ), quackgrass ( Agropyron repens ) in cultivation, and quackgrass in balks were analysed. Two species of eriophyoid mites were recorded: Aceria tosichella Keifer and Abacarus hystrix (Nalepa). A. hystrix appeared more frequent [present in 65.0% of all collected samples; (95% confidence interval: 50–80)] and abundant [mean number of individuals per sample: 207.3 (94.4–351.5)] than A. tosichella (17.5%; (7.5–3.0), 13.8 (0.9–32.8), respectively). Significant differences in numbers among studied mite species populations on different host plants have been found. It is concluded that wheat is the marginal host for these two eriophyoids when compared to quackgrass.  相似文献   

20.
Despite highly specialized pollination strategies, hybridization is a common phenomenon among Mediterranean deceptive orchids. Food-deceptive species sire a progeny of F1 unfertile plants, which work as a late post-zygotic barrier. Conversely, when pre-zygotic barriers of sexually deceptive (Ophrys) species are absent, the hybrids are fertile and an extensive introgression may occur. Here, we have performed molecular analysis and hand pollination treatments to characterize a hybrid zone of two food-deceptive species, O. mascula and O pauciflora. Hybrids (called O. × colemanii) have shown different amounts of parental nrDNA, strongly supporting that they are F2 and/or successive hybrid generations. Comparable high levels of reproductive success have been detected in natural conditions and in experimental crosses suggesting the absence of effective reproductive barriers either between hybrids, either between hybrids and parental species. In light of ecological and distributional features of O. × colemanii across its distribution range, we hypothesize that these populations have originated by secondary contact in the periglacial belt of Apennines. Moreover, the rare and localized O. pauciflora could benefit a genetic enrichment by hybridizing with a widespread related species. O. × colemanii is not a dead end population, but may have a role as potential reserve of adaptive variability and is an unusual stage along the speciation process.  相似文献   

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