Chloroplast DNA markers reveal a geographical divide across Argentinean southern beech Nothofagus nervosa (Phil.) Dim. et Mil. distribution area

  Nothofagus nervosa is one of the most important species of the temperate forests of southern South America. On the eastern slope of the Andes Mountains (Argentina) it has a very small and narrow natural distribution area which follows the valleys of the numerous west-east lake basins. Re-colonisation after the last glaciation is assumed to have originated from refugia located in the western pacific coast or even in free interglacial patches of the Andes Mountains. Sixteen pairs of primers were used to amplify 16 non-coding regions of chloroplast DNA in 11 populations from over the entire distribution area in Argentina. After restriction analysis two polymorphic fragments were found which defined two haplotypes and allowed a differentiation among populations. A clear geographic divide occurs separating populations distributed south and north of a west-east mountain chain. This suggests that after the last glaciation the species spread out from at least two different refugia. Hypotheses about the possible location of these refugia are discussed. Received: 28 October 1997 / Accepted: 19 March 1998     

Colonization routes of Pinus sylvestris inferred from distribution of mitochondrial DNA variation

Understanding the present-day distribution of molecular variation requires knowledge about the history of the species. Past colonization routes and locations of refugia of Scots pine (Pinus sylvestris) were inferred from variation in mitochondrial DNA in material collected from 37 populations located in countries within, and immediately adjacent to the continent of Europe. Two mitochondrial regions, nad1 intron (exon B/C) and nad7 intron 1, were included in the study. Differentiation in maternally inherited mitochondria was high (G _ST′ = 0.824). Two new haplotypes were found at the nad7 intron 1. The occurrence of a 5-bp indel variant was restricted to the Turkish Kalabak population and a 32 bp only found in Central, Eastern, and Northern Europe. The complete absence of the 32-bp indel from the Mediterranean peninsulas supports the view that coniferous forests existed outside these areas during the last glacial maximum, and these populations contributed to the subsequent colonization of the northern parts of Europe. P. sylvestris shares features of its glacial and postglacial history with two other northern, cold-tolerant tree species, Picea abies and Betula sp. These three species differ from many other European trees for which pollen core and molecular evidence indicate colonization from southern refugia after the last glacial period.     

Temporal and spatial diversification of the shrub Justicia areysiana Deflers (Acanthaceae) endemic to the monsoon affected coastal mountains of the southern Arabian Peninsula

Justicia areysiana Deflers (Acanthaceae) is an endemic shrub of the monsoon affected coastal mountains of the southern Arabian Peninsula with ballochorous seed dispersal. Its occurrence in a vegetation belt of relics of the (semi)-evergreen Afro-montane woodland characterised by Olea europaea subsp. cuspidata and Juniperus procera above an elevation of c. 750-800 m is strongly dependent on the activity of the Indian Ocean southwest monsoon. Therefore, the species offers a good model to study the palaeo-climate induced habitat fragmentation of the once continuous distribution range of the Afro-montane woodland belt on the southern Arabian Peninsula. Thirteen populations from the total distribution range of the species were analysed using AFLP fingerprinting and show a distinct geographic distribution pattern with a main split between eastern (Hawf Mountains/Dhofar) and western/central populations (Jabal Urays, Jabal Gedu, Kor Seiban). Our results on the genetic differentiation of populations clearly demonstrate the strong isolation among populations and confirm geographical patterns found in former studies based on chloroplast haplotypes. The dating of the main regional split between eastern and western/central populations based on nrDNA ITS sequence variation, is dated to 0.8–1.8 Myr before present. This indicates drastic habitat fragmentation processes during the Pleistocene which are compatible with periods of aridisation assumed for the African continent at around 1.7 Myr and 1.0 Myr before present.     

Phylogeography and Pleistocene demographic history of the endangered marsh deer (Blastocerus dichotomus) from the Río de la Plata Basin

The marsh deer is the largest neotropical cervid with morphological and ecological adaptations to wetlands and riparian habitats. Historically, this now endangered species occupied habitats along the major river basins in South America, ranging from southern Amazonia into northern Argentina to the Paraná river delta. This particularly close association with wetlands makes marsh deer an excellent species for studying the effects of Pleistocene climatic changes on their demographic and phylogeographic patterns. We examined mitochondrial DNA variation in 127 marsh deer from 4 areas distributed throughout the Río de la Plata basin. We found 17 haplotypes in marsh deer from Brazil, Bolivia and Argentina that differed by 1–8 substitutions in a 601 bp fragment of mitochondrial control region sequence, and 486 bp of cytochrome b revealed only 3 variable sites that defined 4 haplotypes. Phylogeny and distribution of control region haplotypes suggest that populations close to the Pantanal area in central Brazil underwent a rapid population expansion and that this occurred approximately 28,000–25,000 years BP. Paleoclimatic data from this period suggests that there was a dramatic increase for precipitation in the medium latitudes in South America and these conditions may have fostered marsh deer’s population growth.     

The effects of Quaternary glaciations in Patagonia as evidenced by chloroplast DNA phylogeography of Southern beech Nothofagus obliqua

Climatic oscillations during the Quaternary strongly affected the distribution of warm-temperate tree species, which experienced local restrictions into ice-free areas and posterior expansions. To evaluate the impact of these range movements on the genetic structure of populations, we performed a phylogeographical analysis of the species Nothofagus obliqua with chloroplast DNA markers. A total of 27 populations covering the whole natural distribution range were analyzed using polymerase chain reaction-restriction fragment length polymorphism. Diversity (h _T, h _S), allelic richness (r _g), and differentiation among populations for unordered (G _ST) and ordered alleles (N _ST) were calculated. The relationships among haplotypes were evaluated by the construction of a minimum spanning network. The spatial distribution of the genetic variation was analyzed through a Mantel test and with a nested analysis of molecular variance to differentiate between geographic regions. The screening of 11 non-coding regions allowed the identification of 14 haplotypes. A high genetic differentiation was detected (N _ST = 0.875 and G _ST = 0.824) with the existence of phylogeographic structure (p < 0.05). The distribution of the genetic variation was partially explained by the topography of the region when the populations were divided longitudinally into Coastal Mountains, Central Valley, and Andes Mountains (ϕ _RT = 0.093, p = 0.001). In agreement with pollen records, our results support the hypothesis of Coastal refuges since the region harbors high diversity together with older and private haplotypes. Long-lasting persistence of some Coastal populations without contribution to re-colonization is proposed. Additional refuges are also postulated along the Andes and Longitudinal Valley. Survival in multiple glacial refuges is discussed together with the possible migratory routes.     

Postglacial population expansion of Dacrydium pectinatum (Podocarpaceae) in Hainan, southem China, based on cpDNA trnL-F nocoding sequence data

This study determined the sequences of chloroplast DNA (cpDNA) trnL-F non-coding regions of individuals of a tropical coniferous species, Dacrydium pectinatum, collected from 12 natural populations located in Hainan Province, southern China. Sequence length varied from 868 bp to 876 bp, indicating length polymorphism. Base composition in the sequences was high in A+T content between 64.17% and 64.95%, and no recombination event occurred (Rm = 0). Thirty haplotypes were identified based on statistical parsimony algorithm by running the TCS program. Populations of D. pectinatum in Hainan were lacking genetic differentiation. Such a deduction was supported by the observed F _ST values (0.00), AMOVA (24.17% of molecular variance attributed to difference among populations, P>0.05), high values of Nm (ranging from 1.92 to 2.50) and the branching structure in neighbor-joining (NJ) tree constructed from haplotypes. A ‘star-like’ pattern was exhibited in the TCS network of trnL-F haplotypes, and majority of the haplotypes coalesced near the tips in NJ tree. Gene genealogies of cpDNA haplotypes proposed a recent population expansion of D. pectinatum in Hainan, which was further supported by the results from Tajima’s D test and mismatch distribution analysis. Our data, in conjunction with geological and palynological evidences, showed that in the Holocene, due to global warming, refugee populations of D. pectinatum in Hainan might experience a range expansion. __________ Translated from Acta Scientiarum Naturalium Universitatis Sunyatseni, 2005, 44(5): 70–74 [译自: 中山大学学报 (自然科学版), 2005, 44(5): 70–74]     

Isozyme variation and recent biogeographical history of the long-lived conifer Fitzroya cupressoides

Aim Palaeoenvironmental records of Pleistocene glaciation and associated vegetation changes in Patagonia have led to the hypothesis that during the last glacial maximum (LGM) tree species survived locally in favourable habitats. If present populations originated from spread from only one refugium, such as an ice‐free area of coastal Chile (Single Refugium hypothesis), we would expect that eastern populations would be genetically depauperate and highly similar to western populations. In contrast, if the ice cap was not complete and tree species persisted in forest patches on both slopes of the Andes (Multiple Refugia hypothesis), we would expect a greater degree of genetic divergence between populations either on opposite sides of the Cordillera (Cordillera Effect scenario) or towards its present‐day southern distributional limit where the ice sheet reached its maximum coverage (Extent‐of‐the‐Ice scenario). Location We tested this refugia hypothesis using patterns of isozyme variation in populations sampled over the entire modern range of the endemic conifer Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae) in temperate South America. Methods Fresh foliage was collected from twenty‐four populations and analysed by horizontal electrophoresis on starch gels. Results Twenty‐one putative loci were reliably scored and 52% were polymorphic in at least one population. Populations from the eastern slope of the Andes were genetically more variable than those from the western slope; the former had a greater mean number of alleles per locus, a larger total number of alleles and rare alleles, and higher polymorphism. Genetic identities within western populations were greater than within eastern populations. Discriminant analyses using allelic frequencies of different grouping schedules of populations were non significant when testing for the Single Refugium hypothesis whereas significant results were obtained for the Multiple Refugia hypothesis. Main conclusions Our results indicate that present Fitzroya populations are the result of spreading from at least two, but possibly more, glacial refugia located in Coastal Chile and on the southern flanks of the Andes in Argentina.     

Differentiation among Austrian populations of Norway spruce [Picea abies (L.) Karst.] assayed by mitochondrial DNA markers

Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.] across the Austrian Alps and Bohemian Massif were sampled to elucidate the geographical pattern of genetic differentiation. Three polymorphic mitochondrial DNA (mtDNA) loci were surveyed. Two or three alleles were detected at each locus, resulting in seven multilocus mtDNA haplotypes (A–F). Western populations proved to be monomorphic, whereas eastern and central Austrian populations were slightly to highly polymorphic, respectively. As revealed by spatial analysis of molecular variance and Monmonier’s analysis, the two main haplotypes A and B are not randomly distributed. Haplotype A was restricted to central and eastern Austria, whereas haplotype B occurred in all Austrian populations but was the only haplotype identified in western populations. This pattern may be explained by different glacial refugia located in the Dinaric Alps and the Carpathian mountains.     

Phylogeography of Larix sukaczewii Dyl. and Larix sibirica L. inferred from nucleotide variation of nuclear genes

We investigated phylogeography of Larix sukaczewii and Larix sibirica using nucleotide variation at three following nuclear gene regions: 5.8 S rDNA including two internal transcribed spacers (ITS), cinnamyl alcohol dehydrogenase (CAD), and phytochrome-O (PHYO). We also included sequences of the 4-coumarate: coenzyme A ligase (4CL) gene region obtained in our recent study. CAD and PHYO showed very low nucleotide variation, but ITS and 4CL had levels of variation similar to those reported for other conifers. Pleistocene refugia have been hypothesized to exist in the Southern Urals and South Central Siberia, where four out of nine of the investigated populations occur. We found moderate to high levels of population differentiation (F _ST = 0.115 – 0.531) in some pairwise comparisons suggesting limited gene flow and independent evolution of some refugial populations. In L. sukaczewii, low levels of differentiation were found among populations from areas glaciated during the Pleistocene, indicating their recent origin. Our results also suggest these populations were created by migrants from multiple, genetically distinct refugia. Furthermore, some haplotypes observed in populations from previously glaciated areas were not found in putative refugial populations, suggesting these populations might have contributed little to the extant populations created after the Last Glacial Maximum. Some authors regard L. sukaczewii and L. sibirica as a single species, while others consider them as separate species. The observed conspicuous differences in haplotype composition and distribution between L. sukaczewii and L. sibirica, together with high values of F _ST between populations of the two species, appear to support the latter classification. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Ismael A. Khatab and Kariyawasam K.G.U. Hemamali contributed equally to this work.     

Phylogeographic studies on natural populations of the South American fruit fly, <Emphasis Type="Italic">Anastrepha fraterculus</Emphasis> (Diptera: Tephritidae)

Anastrepha fraterculus is an important pest of commercial fruits in South America. The variability observed for morphological and behavioural traits as well as genetic markers suggests that A. fraterculus represents a complex of synmorphic species rather than a single biological species. We studied the correlation between geographical distribution and genetic variation in natural populations from Argentina and south Brazil. Fragments of the mitochondrial gene COII were sequenced in 28 individuals. The matrix of aligned sequences was phylogenetically analysed by parsimony, yielding a cladogram of haplotypes. Based on Templeton’s nested method, no clade showed any geographic pattern for the gene COII, indicating lack of significant association between haplotypic variability and geographic distribution. The analysis of nucleotide substitution distances by Neighbour-Joining algorithm showed that geographically distant populations exhibit low genetic distances. The corresponding trees clustered the populations without showing any geographic pattern. This result suggests that the populations studied are not reproductively isolated.     

Worldwide migration of parasitic mites as a result of bumblebee commercialization

We investigated the status of infestation by a tracheal mite, Locustacarus buchneri, in natural populations of a Japanese native bumblebee species, Bombus hypocrita, collected on Hokkaido Island and in the Aomori prefecture between 1997 and 2001. We also investigated mite infestation in commercial colonies of the European bumblebee, Bombus terrestris, imported from the Netherlands and Belgium, and the Japanese native species, B. ignitus, imported from the Netherlands, between 1997 and 2001. We detected mites in natural populations of the two B. hypocrita subspecies and in the commercial colonies. Analysis of variations in 535 bp sequences of the mitochondrial cytochrome oxidase subunit 1 (CO1) gene showed that the mite haplotypes in the native populations and in the imported colonies did not overlap in 1997–1999, but in 2000–2001 some mites possessing European CO1 haplotypes were detected in the natural populations of Japanese native bumblebees. In addition, many mites possessing Japanese haplotypes were detected in the imported commercial colonies from Europe. Considering the fact that the Japanese native bumblebees, B. hypocrita, were once exported to Europe for commercialization, these results suggest that bumblebee commercialization has caused overseas migration and cross-infestation of parasitic mites among natural and commercial colonies. However, because the Japanese and European CO1 haplotypes were closely related, there was a possibility that the European haplotypes found in the mites in the Hokkaido Island revealed native variation. To clarify the status of mite invasion, further detailed analysis of genetic variation of the mite, using other genetic markers on additional samples, need to be performed.     

祁连山区特有物种黑虎耳草的居群遗传多样性研究

利用叶绿体基因(rbcL和trnS-G)及核糖体DNA内转录间隔区(ITS),对祁连山区狭域分布的特有物种黑虎耳草8个居群(115个个体)进行遗传多样性研究,旨在揭示黑虎耳草的居群遗传结构及其历史进化过程。结果表明:(1)所有取样个体共检测到4个cpDNA单倍型和9个ITS单倍型,其中祁连山东南部的居群固定较多的单倍型和特有单倍型,而西北部居群只固定少数几个广泛分布的单倍型,且遗传多样性普遍较低。(2)基于cpDNA数据和ITS数据的分子变异分析(AMOVA)显示,遗传变异主要来源于居群内。(3)基于cpDNA数据的中性检验表明,Tajima’s D(-1.012 30,P> 0.05)和Fu&Li’s D*(-2.066 77,P> 0.05)均为负值,均不显著;歧点分布分析结果显示,黑虎耳草居群经历过近期的扩张事件。根据物种现有遗传分布格局推测,黑虎耳草在第四纪冰期时退缩到祁连山东南部的边缘避难所,间冰期或冰期后回迁到祁连山西北部地区,在回迁过程中由于奠基者效应导致祁连山西北部的居群仅固定少数广泛分布的单倍型,并呈现出较低的遗传多样性;由于居群较小且相互隔离,该...     

Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum

Aim We examine several hypotheses emerging from biogeographical and fossil records regarding glacial refugia of a southern thermophilic plant species. Specifically, we investigated the glacial history and post‐glacial colonization of a forest understorey species, Trillium cuneatum. We focused on the following questions: (1) Did T. cuneatum survive the Last Glacial Maximum (LGM) in multiple refugia, and (if so) where were they located, and is the modern genetic structure congruent with the fossil record‐based reconstruction of refugia for mesic deciduous forests? (2) What are the post‐glacial colonization patterns in the present geographical range? Location South‐eastern North America. Methods We sampled 45 populations of T. cuneatum throughout its current range. We conducted phylogeographical analyses based on maternally inherited chloroplast DNA (cpDNA haplotypes) and used TCS software to reconstruct intraspecific phylogeny. Results We detected six cpDNA haplotypes, geographically highly structured into non‐overlapping areas. With one exception, none of the populations had mixed haplotype composition. TCS analysis resulted in two intraspecific cpDNA lineages, with one clade subdivided further by shallower diversification. Main conclusions Our investigation revealed that T. cuneatum survived the LGM in multiple refugia, belonging to two (western, eastern) genealogical lineages geographically structured across south‐eastern North America. The western clade is confined to the south‐western corner of T. cuneatum’s modern range along the Lower Mississippi Valley, where fossil records document a major refugium of mesic deciduous forest. For the eastern clade, modern patterns of cpDNA haplotype distribution suggest cryptic vicariance, in the form of forest contractions and subsequent expansions associated with Pleistocene glacial cycles, rather than simple southern survival and subsequent northward colonization. The north–south partitioning of cpDNA haplotypes was unexpected, suggesting that populations of this rather southern thermophilic species may have survived in more northern locations than initially expected based on LGM climate reconstruction, and that the Appalachian Mountains functioned as a barrier to the dispersal of propagules originating in more southern refugia. Furthermore, our results reveal south‐west to north‐east directionality in historical migration through the Valley and Ridge region of north‐west Georgia.     

Genetic variation and phylogeographic history of <Emphasis Type="Italic">Picea likiangensis</Emphasis> revealed by RAPD markers

Repeated cycles of retreat and recolonization during the Quaternary ice ages are thought to have greatly influenced current species distributions and their genetic diversity. It remains unclear how this climatic oscillation has affected the distribution of genetic diversity between populations of wind-pollinated conifers in the Qinghai-Tibetan region. In this study, we investigated the within-species genetic diversity and phylogenetic relationships of Picea likiangensis, a dominant forest species in this region using polymorphic DNA (RAPD) markers. Our results suggest that this species has high overall genetic diversity, with 85.42% of loci being polymorphic and an average expected heterozygosity (H _E) of 0.239. However, there were relatively low levels of polymorphism at population levels and the differences between populations were not significant, with percentages of polymorphic bands (PPB) ranging from 46.88 to 69.76%, Nei’s gene diversity (H _E) from 0.179 to 0.289 and Shannon’s indices (Hpop) from 0.267 to 0.421. In accordance with our proposed hypothesis, a high level of genetic differentiation among populations was detected based on Nei’s genetic diversity (G _ST = 0.256) and AMOVA analysis (Phi _st = 0.236). Gene flow between populations was found to be limited (Nm = 1.4532) and far lower than reported for other conifer species with wide distribution ranges from other regions. No clusters corresponding to three morphological varieties found in the south, north and west, respectively, were detected in either UPGMA or PCO analyses. Our results suggest that this species may have had different refugia during the glacial stages in the southern region and that the northern variety may have multiple origins from these different refugia.     

The potential for seaweed resource development in subarctic Canada; Nunavik,Ungava Bay

Ungava Bay is ice covered 6–7 months of the year and evidence of ice scouring of seaweeds is extensive in the intertidal and shallow subtidal. Maximum tidal amplitudes of 16 m, among the highest in Canadian waters, compound this impact. Despite this level of annual perturbation, very extensive and dense beds of fucoids in the intertidal and laminarians in the subtidal are common on the western shores of Ungava Bay. Ground surveys of 24 intertidal stations combined with satellite images delineated 82,000 tons standing crop of Fucus vesiculosus and Fucus evanescens in Payne Bay, of which 36,000 tons were considered harvestable. Subtidally, kelp cover reached peak biomass at 5–10 m consisting of three primary species, Saccharina longicruris, Laminaria digitata, and Laminaria solidungula. In the area of Payne Bay, kelp beds of 100 ha were common, averaging 9–12 kg m⁻² wet weight. The productivity of brown algae at these latitudes has been assumed to be low relative to southern latitudes. Direct measurement of lineal growth indicates productivity is intermediate between arctic and temperate populations. The potential for medium level industrial harvest exists under conservative management strategies within the constraints of subarctic logistics.     

Molecular phylogeography of Dryas integrifolia: glacial refugia and postglacial recolonization

Chloroplast DNA variation in the Arctic plant species Dryas integrifolia (Rosaceae) was analysed in relation to both the present-day geographical distribution of populations and to Pleistocene fossil records of this species. The phylogeographical structure was weak but the analysis of haplotype diversity revealed several groups of haplotypes having present-day geographical ranges that overlap locations postulated from geographical and fossil evidence to have been glacial refugia. Based on this information we infer that two important refugial sources of Arctic recolonization by this species were Beringia and the High Arctic. Two other putative refugia, located southeast of the ice sheet and along coastal regions of the eastern Arctic may have served as sources for recolonization of smaller portions of the Arctic. The genetic substructure in the species is mostly due to variation among populations regardless of the ecogeographical region in which they are found. Spatial autocorrelation at the regional scale was also detected. High levels of diversity both within populations and ecogeographical regions are probably indicative of population establishment from several sources possibly combined with recent gene flow.     

Intraspecific Genetic Diversity of Undaria pinnatifida in Japan, based on the Mitochondrial cox3 Gene and the ITS1 of nrDNA

The intraspecific genetic diversity of the kelp Undaria pinnatifida (Harvey) Suringar (Laminariales, Phaeophyceae) was investigated using DNA sequences of the mitochondrial cytochrome oxidase subunit 3 (cox3) gene and internal transcribed spacer 1 (ITS1) of nuclear ribosomal DNA in plants collected from 21 localities along the Japanese coast between 2001 and 2003. Morphological variation was also examined and compared with the genetic diversity. Cox3 analyses of 106 plants revealed 9 haplotypes (I–IX) that differed from each other by 1–7 bp (all synonymous substitutions). Haplotype I was distributed in Hokkaido and the northern Pacific coast of Honshu, while haplotype III was found along the Sea of Japan coast of Honshu. Other types were found along the central and southern coast of Honshu. ITS1 analyses of 42 plants revealed 0–1.7% nucleotide differences, but plants from the Sea of Japan coast and northern Japan had similar sequences. The lower genetic differentiation along the Sea of Japan and northern coasts might be due to the recent establishment (after the middle of the last glacial period) of the Sea of Japan flora. The cox3 haplotype of cultivated plants was found in natural populations occurring close to cultivation sites (Naruto, Tokushima Pref., and Hokutan, Hyogo Pref.). This suggests that cultivated plants possibly escaped and spread or crossed with plants of natural populations. Morphological analyses of variation in 10 characters were conducted using 66 plants. The results showed no significant local variation owing to the wide variation in each population and did not support any forma previously described. No correlations between the morphological characters and cox3 haplotypes were detected.     

基于cpDNA trnL-F序列的胡黄连保护遗传学研究

胡黄连为特产中国-喜马拉雅特有高山植物,作为常用中、藏药材,受到灭绝性采挖,作为濒危和二级保护植物亟待科学的保护。该研究以云南和西藏7个野生居群91个个体为材料,基于 cpDNA trnL-F 非编码序列测序分析胡黄连的遗传多样性和遗传结构,分析显著进化单元,确立优先保护居群并提出科学的保护策略。结果表明：胡黄连 trnL-F 序列长度为871~876 bp,根据序列的核苷酸变异共鉴定出5个单倍型,西藏占有2个单倍型,云南占有3个单倍型,西藏和云南2个地区的所有单倍型均不共享。胡黄连具有较低的单倍型多样性(Hd ＝0.43419)和核苷酸多样性(Dij ＝0.00466)。种群间分化度(Fst ＝0.864520)和基因流(Nm ＝0.04)、居群间的遗传分化水平(GST ＝0.916)、AMOVA 分析(0.78％的遗传变异发生在居群内,60.97％的遗传变异发生在地区内居群间,38.25％的遗传变异发生在地区间)均表明,胡黄连居群间存在明显遗传分化。多数一致性树将胡黄连划分为3个进化分支(Ⅰ、Ⅱ、Ⅲ),这3个分支均与地理相关,分支Ⅰ分布于横断山区的4个居群,分支Ⅱ是分布于东喜马拉雅的一个居群,分支Ⅲ是分布于喜马拉雅中段的2个居群。3个分支分属于3个“进化显著单元(ESU)”。这3个 ESU 中白马雪山、茨中、定日、波密、聂拉木五个居群都需要保护,建议现阶段应优先保护的居群是云南白马雪山和西藏波密居群,以就地保护为主。     

Geographic structuring of chloroplast DNA genotypes inTiarella trifoliata (Saxifragaceae)

Tiarella trifoliata comprises varietieslaciniata, trifoliata, andunifoliata, and is distributed from southeastern Alaska to northern California. We analyzed restriction site variation of chloroplast DNA (cpDNA) using 23 endonucleases in 76 populations representing the entire geographic range of the species and the three recognized varieties. We also employed comparative restriction site mapping of PCR-amplified chloroplast DNA fragments using 16 restriction endonucleases. This species exhibits low cpDNA restriction site variation. No differentiation is evident among varieties of this species based on cpDNA data; some plants of each variety were characterized by each of the two major cpDNA types detected. The two major cpDNA clades, which differ by only a single restriction site mutation, are geographically structured. A northern clade comprises populations from Alaska to central Oregon; most populations analyzed from southern Oregon and California form a southern clade. Populations that possess the typical northern cpDNA type also occur disjunctly to the south at high elevations in the Siskiyou—Klamath Mountain area of southern Oregon and northern California. Conversely, the southern cpDNA type is found disjunctly to the north in the Olympic Peninsula of Washington. Both geographic areas characterized by disjunct cytoplasms are considered glacial refugia.Tiarella trifoliata joins two other species,Tolmiea menziesii andTellima grandiflora, in having well-demarcated northern and southern cpDNA lineages. All three species have similar life-history traits and geographic distributions. We suggest that glaciation may have played a major role in the formation of the cpDNA discontinuities present in these three taxa. The pronounced relationship between cpDNA variation and geographic distribution suggests the potential applicability of intraspecific phylogeography to plants via the analysis of intraspecific cpDNA variation. These three examples also join a rapidly growing data base which indicates that cytoplasms are often geographically structured within species and species complexes.