Evidence for multiple origins of Wolbachia infection in springtails

Bacteria of the genus Wolbachia (Rickettsiae) are endosymbionts found in many species of invertebrates. The traditional division of Wolbachia lines into four supergroups has recently been challenged with the finding of a fifth supergroup. Until now, this E supergroup exists of a single host species of the order Collembola (springtails) and is hypothesized to form a ‘missing link’ between Wolbachia lines in higher arthropods and nematodes. We have sequenced the 16S rDNA, the ftsZ and the wsp genes of a Wolbachia lineage present in the sexually reproducing collembolan Orchesella cincta. We tested the hypothesis on the origin of Wolbachia in Collembola by adding this new Wolbachia lineage and a lineage found in the collembolan Mesophorura macrochaeta to the Wolbachia phylogenetic tree. The addition of these two species clearly shows that Wolbachia is not monophyletic in Collembola. We conclude that the origin of Wolbachia in Collembola is a combination of vertical and horizontal transfer between coexisting species.     

甘肃玉米镰孢茎腐病病原菌种群多样性分析

为了解甘肃玉米镰孢茎腐病致病菌的种群结构和数量,于2015和2017年在甘肃省10个市(州)采集玉米茎腐病样品42份,根据形态学特征和EF-1α (tef)基因序列分析进行病原菌的种类鉴定。结果表明,共获得10种镰孢菌,分别为禾谷镰孢菌复合种(Fusarium graminearum species complex, FGSC, 59.3%)、拟轮枝镰孢(F.verticillioides,11.5%)、木贼镰孢(F. equiseti,10.3%)、胶孢镰孢(F. subglutinans,5.9%)、层出镰孢(F. proliferatum, 4.7%)、变红镰孢(F. incarnatum, 4.0%)、三线镰孢(F. tricinctum, 1.9%)、温带镰孢(F. temperatum, 1.2%)、锐顶镰孢(F. acuminatum,0.8%)和尖孢镰孢(F. oxysporum, 0.4%),其中三线镰孢和锐顶镰孢作为玉米茎腐病新病原,属国内外首次发现。禾谷镰孢菌复合种、拟轮枝镰孢、木贼镰孢和胶孢镰孢在甘肃四大生态区(陇东地区、陇南地区、陇中地区和河西走廊)均有分布,其余种仅在1~3个生态区分布。利用镰孢菌的特异性引物EF-1α (tef)对甘肃玉米镰孢茎腐病优势病原菌禾谷镰孢复合种进行种群检测,共鉴定出布氏镰孢(F. boothii)和禾谷镰孢2个种群,其比例为2.75∶1。选用玉米品种甘宇301按照柯赫氏法则进行致病性测定,结果发现10种镰孢菌均可致病。本研究结果为甘肃玉米茎腐病的综合防控提供了科学依据。     

Role of soil mesofauna in dispersal of Coniothyrium minitans: transmission to sclerotia of Sclerotinia sclerotiorum

Coniothyrium minitans Campbell is a mycoparasite with proven biocontrol activity against Sclerotinia sclerotiorum (Lib.) de Bary in the field and glasshouse. It is known to spread from sites of application but the mechanisms of dispersal are unclear. As C. minitans has been recovered from collembolans collected during glasshouse trials, and numerous mites and insects are often associated with decaying S. sclerotiorum-infected plant material in the glasshouse, the mite Acarus siro L. and the collembolan Folsomia candida Willem were used to investigate the potential of soil mesofauna to disperse C. minitans. In an initial investigation, A. siro was found to transmit the mycoparasite from infected to uninfected sclerotia of S. sclerotiorum in moist sterile sand and non-sterile soil. Subsequently, a simple assay system to monitor transfer of C. minitans from colonised wheat grains to uninfected sclerotia of S. sclerotiorum was developed. Both A. siro and F. candida transmitted C. minitans at least 55 mm to sclerotia in soil at water potentials ranging from saturation to −3.6 MPa. Transmission by A. siro was greater in drier conditions (−0.25 to −3.6 MPa) as mites survived poorly in saturated soil. However, water potentials between saturation to −3.6 MPa had no effect on transmission by F. candida, although collembolans died after 18 d at water potentials of −5.4 MPa or drier. Generally, maximum dispersal occurred within 2 weeks. In soil lacking added arthropods, negligible spread of the mycoparasite was observed. These results suggest that soil mesofauna may be important in the dissemination of C. minitans.     

Role of soil mesofauna in dispersal of Coniothyrium minitans: mechanisms of transmission

Soil mesofauna are thought to play a role in the dispersal of Coniothyrium minitans Campbell but the mechanisms of transmission are unclear. The mite Acarus siro L. and the collembolan Folsomia candida Willem are known to be able to transmit the mycoparasite to uninfected sclerotia of Sclerotinia sclerotiorum (Lib.) de Bary in soil, and consequently the mechanisms of transmission of the mycoparasite by these animals have been investigated in detail. To facilitate this work, a novel method for surface sterilising mites was developed. In an initial investigation, A. siro was found to carry inoculum externally and in the gut. Indeed, almost all faecal pellets collected from either A. siro or F. candida, following feeding on C. minitans, contained germinable inoculum of the mycoparasite. Microscopic examination showed that faeces from these animals consisted mainly of conidia or fragments of conidium, but that only intact conidia gave rise to growth of C. minitans. Passage through the gut of either animal reduced percentage germination of conidia to 11 and 60% in mite and collembolan faeces, respectively, compared to 99% in controls not subjected to gut passage. In addition, germination in mite faeces was delayed by approximately 12 h compared to controls. However, subsequent germ tube extension from surviving conidia appeared unaffected by gut passage. Individual faecal pellets of A. siro were found to contain sufficient inoculum of the mycoparasite to initiate infection of sclerotia in moist sterile sand and non-sterile soil. The relevance of these results to the glasshouse and field situation is discussed.     

The nonsusceptibility of soil Collembola to insect pathogens and their potential as scavengers of microbial pesticides

Collembola are very abundant soil insects which contribute to the decomposition of organic matter. This paper presents data on the interaction between Collembola and selected microbial agents being tested or used as biological pesticides. The Collembolan, Folsomia candida, was not susceptible to the entomopathogenic fungi Beauveria bassiana, Metarhizium anisopliae, Hirsutella spp., and Verticillium lecanii, or to the bacterium, Bacillus thuringiensis, and its endotoxins. F. candida consumed and inactivated the insect pathogens without suffering mortality, reproductive disturbance or any other harmful effects. Consumption of insect pathogens demonstrates the potential of the Collembola as scavengers of biological insecticides artificially introduced into the environment.     

MtDNA genetic diversity and population history of a dwindling raptorial bird, the red kite (Milvus milvus)

The red kite (Milvus milvus) occurs in a relatively small area in the southwestern Palearctic region, with population strongholds in Central Europe. Following strong human persecutions at the beginning of the 20th century, populations have receded, particularly in peripheral areas and islands. In order to describe and compare levels of genetic diversity and phylogeographic patterns throughout its entire distribution in Europe, sequence variation of a 357 bps part of the mitochondrial DNA control region was assessed in eight populations and 105 individuals. Overall, results indicate that population declines have affected red kite mtDNA variation. We found low levels of genetic diversity (values of nucleotide diversity ranging from 0 in Majorca island to 0.0062 in Central Europe), with only 10 distinct haplotypes, separated by low levels of genetic divergence (mean sequence divergence = 0.75%). Highest haplotype and nucleotide diversities match with demographic expectations, and were found in Central European and Central Spanish samples, where present strongholds occur, and lowest values in the declining southern Spanish and insular samples. Φst estimates indicated moderate gene flow between populations. Phylogeographic patterns and mismatch distributions analyses suggest central European regions may have been colonized from southern glacial refugia (in the Italian or Iberian peninsulas). Interspecific phylogenetic comparisons and divergence date estimates indicated the genetic split between the red kite and its closely related species, the black kite (Milvus migrans), might be relatively recent. The low level of genetic variation found in the red kite mitochondrial control region, compared to the black kite, is likely the result of relatively recent divergence (associated with founder events), successive bottlenecks and small population sizes. As there are several ongoing projects aimed at reinforcing populations in countries such as the United Kingdom, Italy or Spain, our results may prove useful for the genetic management of the species.     

Genetic differences between the two remaining wild populations of the endangered Indian rhinoceros (Rhinoceros unicornis)

The management of rare and endangered species in the wild and in captivity requires an understanding of the characterization of the genetic units within each species and their relationships to each other. The Indian rhinoceros (Rhinoceros unicornis) is an endangered species with a current population size of c. 2800 individuals. We analyzed 26 individuals of known origin kept in captivity and 21 wild ranging individuals of the two remnant large wild populations in Assam (India) and Nepal employing mitochondrial and microsatellite markers to determine whether the two geographically isolated populations show distinct patterns of genetic diversity, and whether the genetic diversity of the populations is influenced by past demographic bottlenecks. We identified 10 different mitochondrial D-loop haplotypes, of which 4 were specific to the Assam population (10 sequences examined) and 6 specific to the Nepal population (19 sequences). Similarly, the microsatellite analysis demonstrated a strong genetic differentiation between the Assam and Nepal populations and allowed to assign each individual to its origin with high confidence. Furthermore, our analyses revealed the occurrence of a bottleneck in the Assam population long before the reported bottleneck in 1908, and it revealed that the Nepal population is a recent (probably post-glacial) colonization. In summary, the extent of genetic divergence between the two remnant R. unicornis populations suggests separate conservation programs (even for captive individuals) as long as the persistence of the entire species is not severely threatened. The microsatellite markers can also be used to determine the origin of confiscated material such as horns.     

Soil fungal population in preharvest maize ecosystem in different tillage practices in Argentina

The distribution of the soil borne fungal population in maize non-rhizospheric soil in experimental trials were studied during the harvest seasons 2000/2001 and 2001/2002. The effect of different tillage practices was evaluated. Soil samples were collected from three tillage treatments, conventional tillage, reduced tillage and no tillage, with and without grazing. The distribution of fungal propagules in different treatments was compared. Soil samples were collected from the top soil (3 cm) before planting, in the sprong. No tillage without grazing and reduced tillage with grazing had the highest densities of filamentous fungi (5.71 and 5.85 log, respectively). The mycological survey showed the presence of six genera of filamentous fungi. They were Aspergillus, Fusarium, Penicillium, Trichoderma, Cladosporium, Alternaria. The species identified from genus Aspergillus were A. flavus, A. restrictus, A. candidus, A. parasiticus and other Aspergillus spp. The species identified from genus Penicillium were P. pinophylum, P. citreonigrum, P. implicatum, P. purpurogenum, P. minioluteum, P. waksmanii, P. restrictum, P. rugulosum, P. funiculosum and P. variabile. Species from genus Fusarium were identified as F. verticillioides, F. oxysporum, F. proliferatum and others Fusarium species. The isolation frequency of Aspergillus increased in the no tillage and grazing practices treatments. The data indicate that different agricultural practices impact soil inocula of principal maize toxigenic fungi in the production area. To our knowledge, this is the first report on the effect of tillage management practices on fungal populations in preharvest maize agroecosystem in Argentina.     

Current genetic isolation and fragmentation contrasts with historical connectivity in an alpine lizard (Cyclodomorphus praealtus) threatened by climate change

Assessing levels of genetic diversity, connectivity and historical demography for threatened species provides important information for conservation management. We used a combination of the mitochondrial ND4 gene and seven microsatellite markers to examine both historical and recent population genetic structure and demography of the threatened alpine she-oak skink, Cyclodomorphus praealtus. This species is restricted to the “sky islands” of the Australian alpine region. Based on mtDNA, the New South Wales and Victorian regions are reciprocally monophyletic and highly divergent, with among population variation of 0.9 and net sequence divergence of 4.28%, which suggests that they should be considered separate Evolutionary Significant Units for management purposes. The mtDNA data also indicate historical connectivity between the three Victorian populations. However, a model-based clustering analysis of microsatellite genotypes identified strong population structure in Victoria, with three distinct populations that have no current inter-population gene flow. This suggests that the Victorian populations are effectively isolated from each other, and is indicative of very low dispersal capacity and a high degree of habitat specialisation. This is reinforced by the substantially lower genetic diversity within the lowest elevation population compared to the other higher elevation populations. We found no genetic signature of major changes in effective population size. These data provide a baseline for assessing future impacts of climate change on the genetic structure of this alpine endemic species.     

Influence of soil and host plant crop on the genetic diversity of Azospirillum amazonense isolates

The genetic structure of Azospirillum amazonense populations isolated from the rhizosphere soil and washed and surface-sterilised roots of rice, maize and sorghum plants, cropped simultaneously in two different soils (clay loam and sandy loam) was characterised. Genetic diversity was measured by restriction fragment length polymorphism of the amplified 16S–23S rDNA intergenic spacer region (RISA-RFLP) and cluster analysis. Four genetically distinct clusters of isolates were observed with 78% similarity, suggesting that the A. amazonense population was heterogeneous at the strain level regardless of the soil type or host plant. Analysis of molecular variance (AMOVA) demonstrated that the host plant had a highly significant selective effect on the genetic structure of this species, especially on those isolates intimately associated with them, but also to a lesser extent on isolates from the rhizosphere and washed roots. The soil type also had a highly significant selective effect on A. amazonense genetic diversity, especially for those isolates from the rhizosphere soil. The selective effect of the soil type combined with that of the host plant suggests that environmental factors, such as soil texture and composition of exudates provided by C₃ or C₄ plants, play major roles in the overall genetic structure of A. amazonense populations associated with these cereals.     

Spatial distribution of Collembola in presence and absence of a predator

Many Collembola species are characterised by an aggregative behaviour, which is closely related to communication and co-ordination between individuals. In spite of a general heavy predation pressure, a large number of Collembola is found in soil suggesting that they have developed defence strategies against predators.

This study investigates changes in spatial structure of gregarious Collembola species in presence of a predator. It is analysed if the gregarious lifestyle of Collembola populations functions as a defence strategy against predation. In presence of a predator distances between individuals in the Collembola species Folsomia candida and Heteromurus tetrophthalmus became closer, reducing the risk of individuals to be captured by predators. On the contrary, Onychiurus sinensis specimens, which were never caught probably because of the extrusion of repelling fluids by means of their pseudocelli, moved away from each another. Besides being a successful chemical defence against predation, repelling fluid secretion may effectively repel conspecifics, thereby keeping individuals at greater distance.     

Determinants of reproductive success: A comparative study of five endangered river corridor plants in fragmented habitats

River corridor plants in Central Europe have in recent decades become increasingly rare, caused mainly by habitat destruction and change. The aim of this study was to examine the impact of soil quality, but especially of habitat fragmentation in terms of population size and isolation, on seed production and germination of five selected species, all of which being endangered, perennial herbs: Euphorbia palustris, Lathyrus palustris, Pseudolysimachion longifolium, Sanguisorba officinalis, and Senecio paludosus. We sampled totally 58 populations in flood plains in the surroundings of Bremen, North-western Germany.

In all five species, there were positive correlations between the number of seeds (total seed mass) and plant height, as well as between germination and seed mass. Seed traits values and/or germination were negatively affected by small population size, in Euphorbia, Lathyrus and Pseudolysimachion additionally by high population isolation. There were also significant effects of soil quality, but these were less consistent across species. The contribution of population size and isolation, compared to soil quality, to the variance explained in the reproductive components was especially high in Euphorbia and Lathyrus, the only two species depending on insect pollination.

The results indicate that the five species are likely to decline as a consequence of decreased plant fitness in small and/or isolated populations, probably caused by pollen limitation. The studied species may be representative for the group of river corridor plants as a whole that are often restricted to isolated remnants of near-natural flood plain vegetation in a matrix of heavily used agricultural land.     

Microsatellite analysis of genetic population structure of the endangered cyprinid Anaecypris hispanica in Portugal: implications for conservation

The endangered fish species Anaecypris hispanica is restricted to eight disjunct populations in the Portuguese Guadiana drainage. The genetic structure of these populations was studied in order to determine levels of genetic variation within and among populations and suggest implications for conservation of the species. Based on five microsatellite loci, the null hypothesis of population homogeneity was tested. Tests for genetic differentiation revealed highly significant differences for pairwise comparisons between all populations, and substantial overall population subdivision (F_ST=0.112). All sampled populations contained unique alleles. Our findings indicate marked genetic structuring and emphasise limited dispersal ability. The high levels of genetic diversity detected within and among A. hispanica populations suggest, however, that the observed fragmentation and reduction in population size of some populations during the last two decades, has impacted little on levels of genetic variability. Data imply that most A. hispanica populations should be managed as distinct units and that each has a high conservation value containing unique genetic variation. It is argued that geographic patterns of genetic structuring indicate the existence of eight management units.     

Low levels of genetic variation among southern peripheral populations of the threatened herb, Leontice microrhyncha (Berberidaceae) in Korea

Levels of genetic variation and intrapopulation genetic structures of Leontice microrhyncha S. Moore (Berberidaceae) were assessed for six populations in South Korea, representing the southern most range of a species found in Northeast China and the Korean peninsula. Detected genetic diversity (H_es) was very low (0.024) and F_IS values showed large heterozygote deficiencies. The small percentage of polymorphic loci and numbers of alleles per locus suggest that L. microrhyncha has a history of severe or long-lasting population bottlenecks that have eroded genetic diversity. This study suggests that the Korean population appears to consist of two historically isolated and independently evolving populations. It seems likely that these groups have been isolated and unstable for a significant period of time. However, the effects of recent habitat fragmentation on the historically disjunct and fragmented population system found in L. microrhyncha were not those predicted from the lack of significant relationships between population-level patterns of genetic variation and population sizes. Most non-unique genotypes were shared by most individuals and the lower level of diversity, high levels of inbreeding and population differentiation as well as high rate of seed production indicated that this species is autogamous and self-compatible and probably largely selfing. Therefore, to preserve extant genetic variation, all populations must be protected across the small geographic range of the species to retain both allelic and genotypic diversity.     

Conservation genetics of evolutionary lineages of the endangered mountain yellow-legged frog, Rana muscosa (Amphibia: Ranidae), in southern California

Severe population declines led to the listing of southern California Rana muscosa (Ranidae) as endangered in 2002. Nine small populations inhabit watersheds in three isolated mountain ranges, the San Gabriel, San Bernardino and San Jacinto. One population from the Dark Canyon tributary in the San Jacinto Mountains has been used to establish a captive breeding population at the San Diego Zoo Institute for Conservation Research. Because these populations may still be declining, it is critical to gather information on how genetic variation is structured in these populations and what historical inter-population connectivity existed between populations. Additionally, it is not clear whether these populations are rapidly losing genetic diversity due to population bottlenecks. Using mitochondrial and microsatellite data, we examine patterns of genetic variation in southern California and one of the last remaining populations of R. muscosa in the southern Sierra Nevada. We find low levels of genetic variation within each population and evidence of genetic bottlenecks. Additionally, substantial population structure is evident, suggesting a high degree of historical isolation within and between mountain ranges. Based on estimates from a multi-population isolation with migration analysis, these populations diversified during glacial episodes of the Pleistocene, with little gene flow during population divergence. Our data demonstrate that unique evolutionary lineages of R. muscosa occupy each mountain range in southern California and should be managed separately. The captive breeding program at Dark Canyon is promising, although mitigating the loss of neutral genetic diversity relative to the natural population might require additional breeding frogs.     

Genetic structure, population dynamics, and conservation of Black caiman (Melanosuchus niger)

Microsatellite DNA polymorphisms were screened in seven populations of the largest Neotropical predator, the Black caiman Melanosuchus niger (n = 169), originating from Brazil, French Guiana and Ecuador. Eight loci were used, for a total of 62 alleles. The Ecuadorian population had the lowest number of alleles, heterozygosity and gene diversity; populations of the Guianas region exhibited intermediate diversities; highest values were recorded in the two populations of the Amazon and Rio Negro. During the last century Melanosuchus populations have been reduced to 1-10% of their initial levels because of hunting pressure, but no strong loss of genetic diversity was observed. Both the inter-locus g-test and the Pk distribution suggested no recent important recovery and/or expansion of current populations. On a global scale, the inter-population variation of alleles indicated strong differentiation (F_ST = 0.137).Populations were significantly isolated from each other, with rather limited gene flow; however, these gene flow levels are sufficiently high for recolonization processes to effectively act at regional scales. In French Guiana, genetic structuring is observed between populations of two geographically close but ecologically distinct habitats, an estuary and a swamp. Similar divergence is observed in Brazil between geographically proximate “black water” and “white water” populations. As a consequence, the conservation strategy of the Black caiman should include adequate ecosystem management, with strong attention to preservation of habitat integrity. Distribution of genetic diversity suggests that current populations originated from the central Amazonian region. Dispersal of the species may thus have been deeply influenced by major climatic changes during the Holocene/Pleistocene period, when the Amazonian hydrographic networks were altered. Major ecological changes such as glaciations, marine transgressions and a hypothesized presence of an Amazonian Lake could have resulted in extension of Black caiman habitats followed by isolation.     

Mitochondrial DNA sequence, morphology and ecology yield contrasting conservation implications for two threatened buckmoths (Hemileuca: Saturniidae)

Taxa of conservation interest are frequently identified using morphological or ecological characters. These characters are assumed to represent evolutionary importance, population structure and/or phylogenetic relationships in such organisms. We tested this assumption using two species complexes of the moth genus Hemileuca (Saturniidae). Both have populations threatened by habitat loss and need conservation protection. Legislation protects one taxon with apparent ecological differences. We sequenced 624 base pairs of mtDNA from the COI gene for geographically distant populations of the Hemileuca maia species complex and the H. electra species complex. Resultant phylogenies contradict prior assumptions about relationships in both species complexes. The legislatively protected Bog Buckmoth is paraphyletic with widespread H. maia, and its use of a novel hostplant seems to be a local adaptation. Divergent morphology and hostplant use among H. electra subspecies are associated with modest genetic divergence (0.48%). However, a group of unrecognized populations that are morphologically similar and geographically close to H. electra electra have mtDNA that is divergent by an average of 4.1%. There is disagreement regarding prioritization of ecological divergence over neutral genetic distance in conservation. We place ecological variation in a phylogenetic context and recommend that exploration of genetic relationships be undertaken when populations are threatened. Adaptive ecological variation should be evaluated in a phylogenetic context to understand its conservation importance. This study illustrates the importance both of phylogenetic context and the use of independent characters in assessing biodiversity for conservation prioritization.     

Extreme neutral genetic and morphological divergence supports classification of Adriatic three-spined stickleback (Gasterosteus aculeatus) populations as distinct conservation units

The fact that the intraspecific genetic differentiation in neutral genetic markers and genes coding for adaptive traits are not typically correlated has caused a great deal of conceptual and practical trouble in delimitation of conservation units. Although the importance of combining information on adaptive genetic divergence with information on historical and recent gene flow in the delimitation of conservation units has been recognized, integrated empirical studies to this end are still rare. We explored the evidence for the specific conservation status of two freshwater three-spined stickleback (Gasterosteus aculeatus) populations on the Adriatic side of the Balkan Peninsula by comparing their phenotypic and genetic characteristics to those of other representative European populations. Apart from focusing on the neutral genetic divergence in mitochondrial DNA sequences and microsatellite markers, we also compared the patterns of morphological differentiation (i.e. bony armour development) resulting from adaptation to freshwater environments. The Balkanic populations formed two distinct groups with regard to neutral genetic variation and had the least developed bony armour of all the examined populations. All morphometric analyses identified the two Balkanic populations as phenotypically – and hence most likely also ecologically – clearly distinct from other European three-spined stickleback populations. These results suggest that the two Balkanic populations (River Neretva and River Zeta) fulfil the most stringent criteria (i.e. lack of genetic and ecological exchangeability) to be classified as conservation units distinct from other European three-spined stickleback populations.     

Genetics of a butterfly relocation: large, small and introduced populations of the mountain endemic Erebia epiphron silesiana

Species specific levels of genetic diversity are necessary for the viability of populations. Erebia epiphron (Lepidoptera: Satyrinae) is a relic species in the Sudetan Mountains, Czech Republic. These populations represent a suitable study object to test the population genetic effects of large, small and introduced populations by means of allozyme electrophoresis. A large continuous and viable population (>100,000 individuals) shows a significantly higher genetic diversity than a small isolated and declining population (ca. 5000 individuals), which most probably was founded during the Little Ice Age in the 17th and 18th centuries and became isolated afterwards. In 1932 and 1933, 50 females were transferred from the large population to the Krkonose Mountains, where they established a viable and quickly expanding population. The number of transferred individuals was sufficient to transfer most of the allozyme diversity of the large natural population. Due to the higher elevation of the Krkonose in comparison to the native range, this allocation might safeguard the survival of the Czech race of E. epiphron even under conditions of global warming.     

沙打旺种群对草原植被演替影响试验研究

沙打旺人工种群大量进入百里香群落后,促进百里香地椒群落在9～10年短时间内演替到长芒草顶极群落。自然封禁条件下,百里香群落在5～7年较快恢复到百里香+长芒草群落这一偏途顶极,而进一步恢复到顶极长芒草群落则需要很长的时间。沙打旺种群促进这一演替过程的作用机制,在于它迅速将百里香种群几乎完全排挤出群落之外,为长芒草种群占领随着它的自然衰退而空出的空间创造了条件。沙打旺种群引起的这一短期人为演替过程,实际上是原来百里香群落中种群关系重新调整的过程,各个种群数量变化的结局,取决于它们各自的种间竞争能力。一些竞争力比较强的种群,如糙隐子草、兴安胡杖子、阿尔泰狗娃花、大羽茅、厚穗宾草等,由原来百里香群落中的次要伴生种,成为长芒草群落中的主要伴生种,一些竞争力弱的种群,如冷蒿,星毛委陵莱、猪毛蒿、狼毒等,则由原来百里香群落中的主要伴生种,成为长芒草群落中的次要伴生种。