Phylogenetic Relationships of the Silver Saxifrages (Saxifraga, Sect. Ligulatae Haworth): Implications for the Evolution of Substrate Specificity, Life Histories, and Biogeography

The silver saxifrages (Saxifraga sect. Ligulatae Haworth; Saxifragaceae) exhibit remarkable variation of substrate specialization, with strictly calcicole to calcifuge species, as well as life histories which range from semelparity to iteroparity. They occur almost exclusively in the European mountain ranges and display high levels of endemism. Sequences from chloroplast and nuclear ribosomal DNA were obtained to resolve phylogenetic relationships among the silver saxifrages and related taxa and to gain insight into the evolution of substrate specificity, life history, and biogeography. The resulting phylogenies suggested that (1) Saxifraga sect. Ligulatae, as traditionally defined, does not constitute a monophyletic group; (2) lime-secreting hydathodes in calcifuge species apparently represent a secondary nonaptation; (3) semelparity evolved independently two or three times in the silver saxifrages and allied sections, possibly in response to climatic changes that occured during the Pleistocene; and (4) narrow endemics, for example S. cochlearis, likely evolved from the fragmentation of the widespread S. paniculata into refugial populations that became isolated during the glacial maxima of the Pleistocene.     

Trichome anatomy and taxonomy of Saxifraga (Saxifragaceae)

Trichomes on flower pedicels and leaves were examined in 213 species of Saxifraga L. and in the monotypic, segregate genera Zahlbrucknera Reichenb., Saxifragella Engl., Saxifragodes D. M. Moore and Saxifragopsis Small. Pedicel trichomes were found to be better taxonomic characters than foliar ones. Six main trichome types were identified: i) multiseriate, glandular; ii) uniseriate, glandular; iii) sessile, multicellular, glandular; iv) multiseriate, eglandular; v) uniseriate, eglandular; vi) unicellular, eglandular. The different types were distributed (except in certain species) on pedicels as follows. Multiseriate, glandular types occur in four series of Saxifraga sect. Micranthes (Haw.) D. Don, part of sect. Gymnopera D. Don, and in sects Ciliatae Haw., Trachyphyllum Gaudin, Xanthizoon Griseb., Ligulatae Haw., Porophyllum Gaudin, Porphyrion Tausch and Irregulares Haw. They also occur in the genus Saxifragopsis. Uniseriate, glandular types occur in five series of Saxifraga sect. Micranthes , part of sect. Gymnopera and Irregulares , and in sects. Miscopetalum (Haw.) Sternb., Cymbalaria , Griseb. Tridactylites (Haw.) Engl., Saxifraga and Stoloniferae Haw. They also occur in the genus Zahlbrucknera. Eglandular, multicellular trichomes occur occasionally in Saxifraga sects Micranthes and Gymnopera , more frequently in sects Ciliatae and Ligulatae , and are ubiquitous in sects Xanthizoon, Porophyllum and Porphyrion. Eglandular, unicellular trichomes occur only in Saxifragopsis and Saxifragodes. Sessile, glandular trichomes occur in a group of species from Saxifraga sect. Stoloniferae. Saxifragella is glabrous. This taxonomic distribution of trichome types generally supports Engler's classification of the genus, and is compatible with other evidence from palynology, embryology and seed morphology. The taxonomic disposition of the few exceptional species is discussed.     

Possible influences of habitat characteristics on the evolution of semelparity and cannibalism in the hump earwigAnechura harmandi

The possible influences of life history and habitat characteristics on the evolution of semelparity and cannibalism in the hump earwigAnechura harmandi were studied. This species is univoltine and overwinters as an adult. Females laid single egg-batches during winter in nests under stones at a riverside in a valley. They took care of the eggs which hatched in early spring and the offspring ate their mother before dispersing. The valley was sometimes flooded in summer. Nymphs emerged as adults and dispersed to elsewhere before the rainy season arrived. They returned to the riverside after the rainy season. The flooding and/or summer heat seemed to be the selective force for the evolution of dispersal behavior and semelparity in this species. The cannibalism of the female parent by her offspring seemed to have readily evolved after the evolution of semelparity. The unfavorable environmental conditions seemed to have a large effect on the evolution of semelparity and cannibalism in this species.     

Circumscription and phylogeny of Saxifraga sect. Ciliatae: Evidence from nrDNA ITS sequences

The largest section of the genus Saxifraga (Saxifragaceae), sect. Ciliatae, consists of 175 morphologically diverse species. This section is mainly distributed in the Qinghai-Tibetan Plateau and adjacent regions of southwest China and more than 80% of the total number of species are endemic to this region. It remains unknown whether this section is monophyletic and up to now no study has been conducted on the infra-sectional phylogeny. In this study, ITS sequences of the nuclear ribosomal DNA were firstly determined for 33 species mainly from this section and related sections. We further downloaded the corresponding sequences of the same DNA region for the other 22 species of Saxifraga and Mitella from GenBank. All sequences were together used to construct the phy-logenetic trees. The main implications of the phylogenetic analyses include: (1) sect. Ciliatae, as traditionally defined, constitutes as a monophyletic clade and its sister group is a well supported clade that includes species from 8 sections such as sect. Porphyrion, sect. Saxifraga and sect. Mesogyne; (2) three morphological subsections, i.e., subsect. Gemmiparae, subsect. Hirculoideae and subsect. Rosulares were tentatively recovered despite the relatively low statistic bootstrap support for the last one; however, subsect. Flagellares and subsect. Hemi-sphaericae were not recognized as separate entities, and nested within subsect. Gemmiparae; (3) subsect. Hircu-loideae and subsect. Rosulares clustered together as sister subclades while subsect. Gemmiparae diverged early. In addition, our results suggest that the paired variation of ITS sequences in sect. Ciliatae is relatively low between the sampled species in spite of their diverse morphology. It is suggested that such a scenario may mirror rapid speciation in this section that probably trigged by the uplifts of the Qinghai-Tibetan Plateau and the extensive selection pressure under the alpine environments.     

Reproductive strategies in some arctic Saxifraga (Saxifragaceae), with emphasis on the narrow endemic S. svalbardensis and its parental species

Arctic saxifrages show conspicuous reproductive and chromosomal variation. We examined sexual and asexual traits in 43 phytotron-cultivated Svalbard populations of nine species, including the endemic, supposedly entirely asexual and aneupolyploid S. svalbardensis and its parental species, S. cernua and S. rivularis. All species were self-compatible hermaphrodites with low pollen/ovule ratios, including the strongly protandrous S. cernua , which previously has been reported as self-incompatible with an androdioecious mating system. Spontaneous selfing resulted in considerable seed set in several species and a few seeds in S. svalbardensis and S. cernua; hand-selfing and cross-pollination often increased seed set in the two latter species. Self-fertilized seeds of S. svalbardensis and S. cernua were viable and developed into normal, vigorous plants. Saxifraga rivularis and its close relative S. hyperborea were strongly autogamous. The bulbil-reproducing S. svalbardensis and S. cernua showed extreme variation in fertility, probably because of frequent aneuploidy. Many plants of S. cernua were fully fertile, suggesting that although natural seed set rarely has been observed, sexual reproduction is frequent enough to maintain its previously reported high levels of clonal diversity. Some plants of S. svalbardensis were also fairly fertile. This species may have considerable evolutionary potential; sexual events can lead to increasingly fertile genets with euploid chromosome numbers.     

Boom and bust: a review of the physiology of the marsupial genus <Emphasis Type="Italic">Antechinus</Emphasis>

The marsupial genus Antechinus is a group of small carnivorous marsupials from the order Dasyuromorphia (Family Dasyuridae) and is found in eastern Australia. The life history of all species in the genus is characterized by a complex, but highly synchronized life cycle in both sexes, culminating in a short mating period followed by total male mortality (semelparity). The breeding season is defined by a specific rate of increase in photoperiod, which is different for each species. In Antechinus spp., male mortality is due to the effects of high free testosterone and cortisol levels on many organ systems. Unusually, spermatogenesis is complete before testosterone levels begin to rise at the winter solstice. In males, low sperm counts have been compensated for by high proportions of sperm reaching the isthmus of the female reproductive tract and long-term storage in the crypts. The females survive to rear their young and may mate again in their second year. Gestation lasts from 26 to 34 days, depending on the species. However, developmental arrest can occur at several stages during embryogenesis, elongating the apparent gestation duration by several days. Several species have strong female sex biases in their litters. The high degree of life history synchrony and the cascade of endocrine-driven physiological events that result in male death are unusual physiological characteristics for mammals. Suggestions why semelparity may have evolved in Antechinus are discussed.     

Molecular analysis of the Pleistocene history of Saxifraga oppositifolia in the Alps

A recent circumpolar survey of chloroplast DNA (cpDNA) haplotypes identified Pleistocene glacial refugia for the Arctic-Alpine Saxifraga oppositifolia in the Arctic and, potentially, at more southern latitudes. However, evidence for glacial refugia within the ice sheet covering northern Europe during the last glacial period was not detected either with cpDNA or in another study of S. oppositifolia that surveyed random amplified polymorphic DNA (RAPD) variation. If any genotypes survived in such refugia, they must have been swamped by massive postglacial immigration of periglacial genotypes. The present study tested whether it is possible to reconstruct the Pleistocene history of S. oppositifolia in the European Alps using molecular methods. Restriction fragment length polymorphism (RFLP) analysis of cpDNA of S. oppositifolia, partly sampled from potential nunatak areas, detected two common European haplotypes throughout the Alps, while three populations harboured two additional, rare haplotypes. RAPD analysis confirmed the results of former studies on S. oppositifolia; high within, but low among population genetic variation and no particular geographical patterning. Some Alpine populations were not perfectly nested in this common gene pool and contained private RAPD markers, high molecular variance or rare cpDNA haplotypes, indicating that the species could possibly have survived on ice-free mountain tops (nunataks) in some parts of the Alps during the last glaciation. However, the overall lack of a geographical genetic pattern suggests that there was massive immigration of cpDNA and RAPD genotypes by seed and pollen flow during postglacial times. Thus, the glacial history of S. oppositifolia in the Alps appears to resemble closely that suggested previously for the species in northern Europe.     

On the origin of the Norwegian lemming

The Pleistocene glacial cycles resulted in significant changes in species distributions, and it has been discussed whether this caused increased rates of population divergence and speciation. One species that is likely to have evolved during the Pleistocene is the Norwegian lemming (Lemmus lemmus). However, the origin of this species, both in terms of when and from what ancestral taxon it evolved, has been difficult to ascertain. Here, we use ancient DNA recovered from lemming remains from a series of Late Pleistocene and Holocene sites to explore the species' evolutionary history. The results revealed considerable genetic differentiation between glacial and contemporary samples. Moreover, the analyses provided strong support for a divergence time prior to the Last Glacial Maximum (LGM), therefore likely ruling out a postglacial colonization of Scandinavia. Consequently, it appears that the Norwegian lemming evolved from a small population that survived the LGM in an ice‐free Scandinavian refugium.     

Foliar crystals in Saxifraga and segregate genera (Saxifragaceae)

In the genera Saxifraga, Saxifragodes, Saxifragella, Saxifragopsis and Zahlbrucknera foliar crystals were found to be present only in Saxifragodes and Saxifraga sections Micranthes and Irregulares . The crystals occur as stellate druses, except in Saxifraga nipponica and S. cortusifolia (section Irregulares ) where they occur as single needles. This finding indicates a close relationship between these two sections of Saxifraga . The monotypic Saxifragodes , from southern South America, does not appear to be related to Andean species of Saxifraga , which lack crystals, but rather to species of Saxifraga section Micranthes in North America. The taxonomic affinities of five species in section Micranthes that lack crystals are discussed.     

The relationship of Sophora sect. Edwardsia (Fabaceae) to Sophora tomentosa, the type species of the genus Sophora, observed from DNA sequence data and morphological characters

Sophora tomentosa , the type species of the genus Sophora , is shown by phylogenetic analyses of rbc L and ITS sequence data to be sister to Sophora sect. Edwardsia . S. tomentosa and most of the species from sect. Edwardsia share hypogeal germination, exstipulate leaves, and terete filaments. These species have buoyant seeds, and are distributed by ocean currents throughout the pantropics ( S. tomentosa ) and around southern temperate oceanic islands (sect. Edwardsia ). S. tomentosa differs from the species of sect. Edwardsia by its frutescent growth habit, terminal elongate inflorescence and smooth-walled legume. S. macrocarpa is unusual in sect. Edwardsia as its leaves have stipules, the filaments are winged, and the legume is smooth-walled.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 439–446.     

Differences between calcifuge and acidifuge plants in root exudation of low-molecular organic acids

The nature and quantity of low-molecular organic acids (LOAs) exuded by the roots of nine species of calcifuge and nine species of acidifuge wild plants from northern Europe were determined by ion chromatography. Particular attention was paid to differences between the calcifuge and the acidifuge species in the proportions of different LOAs in their root exudates. Great differences in mol% root exudation between the calcifuge and the acidifuge species were found in some acids. The calcifuge species exuded more acetic acid, the acidifuge species more oxalic acid and much more citric acid. In three calcifuge species, however, root exudation of oxalic acid was appreciable, whereas acetic acid exudation was low in these species. The phosphate- and Fe-solubilizing ability of eight LOAs in a rhizosphere limestone soil was also tested. Oxalic acid was the most efficient phosphate solubilizer and citric acid, by far, the most efficient Fe-solubilizer at the concentration (10 mM) tested. It might be hypothesized that acidifuge species use oxalate to solubilize phosphate and citrate to solubilize Fe, in limestone soil. The inability of calcifuge species to grow in limestone soil might, therefore, be due to low root exudation of these acids and, as a result, inability to solubilize phosphate and Fe in limestone soil.     

Maternal Care, Iteroparity and the Evolution of Social Behavior: A Critique of the Semelparity Hypothesis

The Semelparity Hypothesis (Tallamy and Brown in Animal Behav 57:727–730, 1999) predicts that among insects with parental care that iteroparity will be rare. It represents two important challenges. First, life history ecologists have sometimes linked extended parental care with iteroparity, not semelparity, as part of a suite of correlated characters associated with K-selective environments. Second, behavioral ecologists have developed theories for the evolution of eusociality that rely upon a subsocial species producing multiple cohorts of offspring, a precondition for offspring allocare and/or inheritance of a social unit. Using a database of invertebrates exhibiting maternal care in Costa (The other insect societies. Harvard University Press, Cambridge, 2006), the association between semelparity and maternal care was tested using a broad comparative analysis. Semelparity was found in only 24.5 % of the best-studied representative species. In addition, semelparity was more rare in species that form nests, burrows or galleries (12.1 %) than in species that guard offspring out in the open (45.0 %). Iteroparity was common both among nesting species with non-overlapping broods (serial nesting) and in species where a female produces broods of different aged offspring in the same nest (within-nest iteroparity). It is hypothesized that common factors, particularly rapid juvenile development on high quality resources, facilitated both serial nesting and parental care. Within-nest iteroparity is an essential stage in the evolution of eusociality that has often been overlooked. Recent models of sibling conflict and reproductive spacing suggest that parental care can be an indirect cause of within-nest iteroparity despite the fact that parental investment can lead directly to diminished future reproduction. The reversal of this life history correlation may occur as a result of the transition between asocial and subsocial nesting behavior; analogous reversals may be a frequent outcome of transitions between levels of social organization.     

中国虎耳草属(虎耳草科)二新种————班玛虎耳草和丁青虎耳草

记载了中国虎耳草属Saxifraga二新种, 即班玛虎耳草S. banmaensis J. T. Pan和丁青虎耳草S. dingqingensis J. T. Pan。其中, 班玛虎耳草仅见于青海班玛, 与小伞虎耳草S. umbellulata Hook. f. &; Thoms.近缘, 但其萼片先端具软骨质短尖头, 花瓣线形, 非提琴状长圆形至提琴形, 基部无爪, 可资区别。丁青虎耳草见于西藏丁青, 与近加拉虎耳草S. llonakhensis W. W. Smith相似, 但其萼片3脉, 于先端汇合成1疣点, 花瓣具8痂体和4-5脉, 基部截形或近耳形, 可以区别。此两种均系中国特有种, 隶属于莲座状亚组subsect. Rosulares Gornall。     

Iron and phosphate uptake explains the calcifuge–calcicole behavior of the terricolous lichens Cladonia furcata subsp. furcata and C. rangiformis

Mechanisms causing the calcifuge–calcicole behavior of lichens are largely unexplored. Studying the case examples of two closely related terricolous lichens, the calcifuge Cladonia furcata subsp. furcata and the calcicole C. rangiformis, we found that preference for acidic or calcareous soils in these lichens is related to iron and phosphate uptake as in vascular plants. In laboratory studies, the calcicole species was more efficient in the intracellular uptake of Fe³⁺ and phosphate at pH 8 than the calcifuge species. At pH 3, intracellular uptake of Fe²⁺ in the calcicole species significantly exceeded that in the calcifuge species suggesting that calcicole lichens suffer from toxicity symptoms by excess Fe²⁺ at acidic sites. Though these observations parallel findings from calcifuge and calcicole vascular plants, mechanisms leading to the different iron and phosphate uptake characteristics in the studied calcifuge and calcicole lichens may differ from those in vascular plants and should be the topic of future research. A role of the depside atranorin in facilitating iron uptake by reducing Fe³⁺ in the apoplast is hypothesized.     

Evolution of Synthyris sect. Dissecta (Plantaginaceae) on sky islands in the Northern Rocky Mountains

We examined the phylogeography of Synthyris sect. Dissecta (Plantaginaceae), which is restricted to the Olympic (S. lanuginosa) and Northern Rocky Mountains (S. dissecta and S. canbyi), to infer effects of sky islands and modes of speciation. Sequences of cpDNA trnT-trnL and psbA-trnH intergenic spacers resolved 22 haplotypes among 185 individuals sampled from 16 populations of the three species. Gene flow in the ancestral lineage and random capture of haplotypes in species lineages of sect. Dissecta have resulted in haplotype clades that are not exclusive to species. Nested clade analysis (NCA) indicates that allopatric fragmentation separated Olympic and Northern Rocky Mountain populations, giving rise to the Olympic endemic S. lanuginosa, which is characterized by unique haplotypes consistent with long temporal isolation. Low haplotype and nucleotide diversity in S. canbyi are consistent with newly founded populations experiencing a bottleneck. Furthermore, we infer S. canbyi evolved as a peripheral isolate of S. dissecta. NCA indicated limited migration in S. dissecta with possible isolation by distance. Both isolation on interglacial sky islands and valley glaciers during at least the last glaciation limited gene flow among populations of S. dissecta in different ranges of the Northern Rocky Mountains.     

A phylogenetic study ofSaxifraga sect.Saxifraga (Saxifragaceae) based on nrDNA ITS sequences

Parsimony analyses of 54 nrDNA ITS (Internal Transcribed Spacer) sequences ofSaxifraga sect.Saxifraga were performed. In addition to some unresolved clades, there is strong disagreement between the ITS phylogeny and previous classifications based primarily on morphology. The extensive cytological instability of sect.Saxifraga prevents previous cytotaxonomical results from resolving the incongruence between molecular and morphological data. Dissimilar topologies between chloroplast (matK) and nuclear (ITS) trees for eight species of sect.Saxifraga suggest that gene trees and the true species tree are not coincident. Recent and mid-term reticulation is proposed as an explanation for the incongruence between morphological, cytological, organellar, and nuclear data. Homogenization in multigene families, such as the ITS region, via concerted evolution may be the key to the interpretation of results based on ITS sequences within sect.Saxifraga. The use of organellar genes in a larger sample should help to determine whether extensive reticulation occurs in sect.Saxifraga, as has been documented in various genera of Saxifragaceae.     

Phylogeographic history of the land snail Candidula unifasciata (Helicellinae,Stylommatophora): fragmentation,corridor migration,and secondary contact

Abstract.— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.     

中国新生代晚期的剑齿象(剑齿象科,长鼻目)及其扩散事件

对我国新生代晚期的剑齿象(Stegodon)进行再研究,提出了下列几点看法:1)依据Ste-godon的特征,可将已命名的中国剑齿象种分为3类。一类为有效的剑齿象种,包括晚中新世的S.licenti Teilhard & Trasseart,1937;上新世的S.zdanskyi Hopwood,1935和S.officinalis Hopwood,1935;上新世晚期至更新世早期的S.zhaotongensis Chow & Zhai,1962;更新世的S.chiai Chow & Zhai,1962,S.orientalis Owen,1870,S.sinensis Owen,1870和S.huananensis sp.n.等种。另一类是从剑齿象属中分出,归入到其他属或科的种。属于这一类的有可能置入真象类(Elephantoidae)的Stegodon parahypsilophus He,1984,S.guizhouensis Li & Wen,1986,Stegodon cf.S.hypsilophus和S.primitium Liu et al.,1973以及归入Stegolophodon的Ste-godon baoshanensis Yun,1975。第三类为属种分类位置未定的种,以S.wushanensis Huang et al.,1991为代表。2)南亚更新世的S.elephantoides(Clift),1828可能在我国不存在。3)我国的剑齿象化石相当丰富,发现于东经99°以东、北纬38°以南的广大地区(包括山西、陕西和甘肃以及华中、华南、西南和部分华东等地区),生存时代为晚中新世晚期至更新世。4)在新生代晚期我国的剑齿象可能经历了3次大的由北向南的扩散和迁移事件。第一次发生在晚中新世,第二次在晚中新世晚期至早上新世早期,第三次在更新世早期。5)剑齿象可能起源于亚洲。S.licenti是我国出现时代最早的剑齿象,也可能是剑齿象最原始的一种。     

The Origin,Dispersal and Formation of the Distribution Pattern of Swertia L. (Gentianaceae)

The genus Swertia is one of the large genera in Gentianaceae, including 154 species, 16 series and 11 sections. It is disjunctly distributed in Europe, Asia, Africa and N. America, but entirely absent from Oceania and S. America. According to Takhtajan’s (1978) regionalization of the world flora, Swertia is found in 14 regions. Eastern Asiatic region with 86 species, of which 58 are local endemics, 13 series and 9 sections, ranks the first among all the regions. The highest concentration of the taxa and endemics in Eastern Asiatic region occurs in SW China-Himalayan area (Sikang-Yunnan P. , W. Sichuan, W. Yunnan-Guichou Plateau of China and NE. Burma, N. Burmense P. , E. Himalayan P. and Khasi-Manipur P. ). In this area there are 74 species (48 endemics), 12 series, and 9 sections; thus about half species of the world total, three quarters of series and 82% of sections occur in this small area. Besides, the taxa at different evolutionary stages in Swertia also survive here. It is an indication that SW. China-Himalayan area is a major distribution centre of the genus Swertia. In addition, Sudan-Zambezian Region in Africa, with 22 species, 4 series and 2 sections, is a second distribution centre. The primitive type of the genus Swertia is Sect. Rugosa which consists of 2 series and 23 species. It is highly centred in the mountains of SW. China (Yunnan, Sichuan, Guizhou and SE. Xizang) where 2 series and 16 species occur. Among them 15 species of Ser. Rugosae were considered as the most primitive groups in this genus. From our study, the outgroup of Swertia is the genus Latouchea Frahch. , which is distributed in Yunnan, Sichuan, Guizhou, Hunan, Guangdong, Guangxi and Fujian. The two groups overlap in distribution in SW. China. According to the principle of common origin, the ancestor of two genera ap peared most probably in this overlapping area. It was inferred that SW. China Was the birth-place of the genus Swertia. Four sections of Swertia have different disjunct distribution patterns: Sect. Ophelia is of Tropic Asia, Africa and Madagascar disjunct distribution; sect. Swertia is of north temperate distribution; sect. Spinosisemina is in Tropical Asia (Trop. India to S. China and Philipines); sect. Platynema also is in Tropical Asia (Java, Sumatra, Himalayas to SW. China). These disjunct patterns indicate that the Swertia floras between the continents or between continent and islands have a connection with each other. From paleogeographical analysis, Swertia plants dispersed to Madagascar before the Late Cretaceous, to SE. Asian Islands in the Pleistocene, to North America in the Miocene. The distribution of Swertia in Madagascar might be later than that in Asia. Therefore the origin time of the genus Swertia was at least not later than the Late Cretaceous, and might be back to the Mid-Cretaceous. The genus Swertia first fully developed and differentiated, forming some taxa at different evolutionary stages (Rugosa, Swertia, Poephila, Ophelia and Platynema etc. ) in the original area, and these taxa quickly dispersed in certain directions during the Late Cretaceous-Middle Tertiary when the global climate was warm and no much change. There seem to be three main dispersal routes from the origin area to different continents; (1) The westward route i. e. from SW. China, along the Himalayas area to Kashmir, Pakistan, Afghanistan and Iran, and then southwestwards into Africa throuth Arabia. Four sections (Poephila, Macranthos, Kingdon-Wardia and Ophelia) took this dispersal route. Most species of sect. Ophelia dispersed along this route, but a few along southern route and north ern route. Sect. Ophelia greatly differentiated in Africa and the African endemic sectionSect. Montana was derived from it. The two sections form there a second distribution center of Swertia. (2) The southward route, i. e. towards S. India through the Himalayas, and towards SE. Asian islands through C. and S. China, Indo-China. Along this dispersal route sect. Platynema, Sect. Spinosisemina and a few species of Sect. Ophelia dispersed; (3) The northward rout, i. e. northwards across N. China, C. Asia to a high latitude of Euasia, and also through E. Asia into N. America. The following groups took this route: sect. Rugosa, sect. Swertia, sect. Frasera, sect. Heteranthos and sect. Ophelia ser. Dichotomae. Therefore, it seems that the genus Swertia originated in SW. China and then dispersed from there to N. and S. Asia, Africa, Europe and North America and formed the moderndistribution pattern of this genus.     

Some ecophysiological and historical approaches to species richness and calcicole/calcifuge behaviour — contribution to a debate

Species richness in vascular plants was related to the plants’ calcifuge or calcicole behaviour using documentation from forests and open-land vegetation at about one thousand sites in the southern parts of Sweden. It is concluded that vegetation of strongly acid soils (pH-KCl < 4.5) have fewer vascular plant species than comparable vegetation of other soils, whereas there are no consistent differences in species richness between slightly-moderately acid and calcareous sites. Low species richness is particularly related to high concentrations of Al³⁺ and H⁺ ions (either soil solution concentrations or concentrations of exchangeable ions), not to a lack of calcium carbonate. The majority of plant species are able to render the sparingly soluble phosphate, iron and manganese compounds of high-pH soils available, but they are unable to tolerate much Al³⁺ or H⁺. Acidicole (calcifuge) species have developed the power of tolerating Al³⁺ and H⁺, which may be considered a secondary property of plants, but they have lost the power of solubilizing critical mineral nutrients in high-pH soils. The reasons why these ecophysiological properties are inversely related in the current flora are obscure, difficult to account for experimentally and a main ecological problem. In areas with cool-temperate climates the flora was partly or mainly extinguished by the Pleistocene glaciations. Comparatively fewer calcifuge than calcicole species have, since then, had enough time to develop, and the number of calcifuges is lower, in spite of the fact that most natural and seminatural soils of these areas are nowadays acidic.