Biogeographic patterns in Mediterranean and Macaronesian species of Saxifraga (Saxifragaceae) inferred from phylogenetic analyses of ITS sequences

A biogeographic study of Saxifraga section Saxifraga was performed based on phylogenetic analyses of ITS (internal transcribed spacer) sequences of nuclear ribosomal DNA. ITS sequences from 21 species and 31 populations were examined to identify colonization patterns for the two species of Saxifraga occurring in Macaronesia and for S. globulifera in the west Mediterranean basin. Phylogenetic analysis of the sequence data yield a single most parsimonious tree with many of the major clades well supported by bootstrap and decay values. The ITS tree provided resolution at specific and populational levels that points to two biogeographic patterns within the genus. In contrast to the molecular evidence provided by other authors for a Mediterranean origin of several Macaronesian genera of angiosperms, our results indicate that the Madeiran archipelago was colonized a single time by a species of Saxifraga originating from the Eurosiberian region. On the other hand, the molecular evidence also suggests that populations of S. globulifera from North Africa have been isolated for a long time from populations occurring in the Iberian Peninsula, and that the endemic S. reuteriana has evolved from the Iberian populations of S. globulifera. The Mediterranean Sea has probably been an effective isolating barrier for some plant groups that occur in Europe and North Africa.     

Chloroplast phylogenomics and the taxonomy of Saxifraga section Ciliatae (Saxifragaceae)

Comprising ca. 200 species, Saxifraga sect. Ciliatae is the most species-rich section of Saxifraga s.str., whose center of diversity is in the Tibeto-Himalayan region. The infra-sectional classification of sect. Ciliatae is still in debate due to the high level of species richness, as well as remarkable variations of habitat, morphology, physiology and life cycles. Subdivisions of this section proposed in various taxonomic systems have not been adequately tested in previous phylogenetic studies, partly due to low taxonomic sampling density, but also to the use of few DNA markers. In order to achieve a more robust infra-sectional classification of sect. Ciliatae, complete chloroplast genomes of 94 taxa from this section were analyzed, of which 93 were newly sequenced, assembled and annotated. The length of the 94 plastomes of sect. Ciliatae taxa range from 143,479 to 159,938 bp, encoding 75 to 79 unique protein-coding genes (PCGs). Analyses of the 94 plastomes revealed high conservation in structural organization, gene arrangement, and gene content. Gene loss and changes of IR boundaries were detected but in extremely low frequency. The molecular phylogenetic tree from concatenated PCGs and complete chloroplast genome sequences exhibited high resolution and support values and confirms that sect. Ciliatae is monophyletic. Three well-supported clades were revealed within the section that agree relatively well with the subsectional taxonomy of Gornall (1987), but some minor modifications should be made. Firstly, the monotypic subsection Cinerascentes should be abandoned and its constituent species, S. cinerascens, assigned to subsect. Gemmiparae. Secondly, subsections Rosulares and Serpyllifoliae should be merged and become subsect. Rosulares. Section Ciliatae thus comprises: subsect. Hirculoideae Engl. & Irmsch.; subsect. Rosulares Gornall; subsect. Gemmiparae Engl. & Irmsch.; subsect. Flagellares (C. B. Clarke) Engl. & Irmsch. and subsect. Hemisphaericae (Engl. & Irmsch.) Gornall.     

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。     

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.     

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.     

Understanding population history for conservation purposes: population genetics of Saxifraga aizoides (Saxifragaceae) in the lowlands and lower mountains north of the Alps

Several alpine species have outlying populations in the lowlands and lower mountains north of the Alps. These small, isolated populations are usually described as either (1) glacial relics, (2) descendants from populations living on forelands and moraines during the ice ages, or (3) populations founded by long-distance dispersal after glaciation. A floristic survey of the historic and present distributions and an allozyme investigation were performed on one of these relic species, Saxifraga aizoides. The species was historically more abundant and had more stations in more regions of northeastern Switzerland. The former population structures within regions, nowadays destroyed, were still reflected in distinct and high regional genetic diversity and variation. There was weak evidence of increased inbreeding in outlying populations, but populations did not deviate from Hardy-Weinberg equilibrium. No geographic pattern of genetic variation above the regional scale (>10 km) was found. Based on the spatial and genetic structures found, it was not possible to discriminate between the abovementioned hypotheses. Nevertheless, the study shows how a thorough evaluation of distribution and abundance data aids the interpretation of genetic data with respect to population history, biogeography, and conservation biology.     

matK and rbcL gene sequence data indicate that Saxifraga (Saxifragaceae) is polyphyletic

The large genus Saxifraga, which consists of ≈400 morphologically and cytologically diverse species, has long been considered taxonomically complex. Phylogenetic analysis of over 2500 bp of chloroplast sequence data derived from matK and rbcL was employed to examine relationships among sections of Saxifraga, the segregate genera Zahlbrucknera, Saxifragopsis, and Cascadia, and the relationships of these taxa to other Saxifragaceae sensu stricto. Phylogenetic trees resulting from separate analyses of the matK and rbcL sequences were highly congruent; phylogenetic analysis of a combined matK–rbcL data matrix was therefore also conducted. Our analyses indicate that Saxifraga is polyphyletic, comprising two well-differentiated clades. One clade, Saxifraga sensu stricto, is the sister to the remainder of the family and consists of Saxifraga sections Irregulares, Heterisia, Trachyphyllum, Cymbalaria, Mesogyne, Xanthizoon, Porphyrion, Ciliatae, Cotylea, Ligulatae, Saxifraga, and Gymnopera. With the exception of Gymnopera, the species-rich sections of this clade are monophyletic. Also part of this clade is the problematic Zahlbrucknera paradoxa, which is allied with members of section Saxifraga. A second major clade of Saxifraga species, Micranthes sensu lato, comprises the large section Micranthes, as well as the segregate genus Cascadia, and S. tolmiei of section Merkianae. This clade is allied with the Heuchera, Darmera, and Chrysosplenium-Peltoboykinia groups of genera. The segregate genus Saxifragopsis is only distantly related to species of Saxifraga, and is instead the sister to Astilbe. The monotypic Oresitrophe is confirmed as a member of the Darmera group of genera. These results suggest that the floral features used to define Saxifraga may simply be symplesiomorphic in these well-separated Saxifraga lineages. Furthermore, the enormous cytological diversity encompassed by Saxifraga likely represents two independent instances of extensive aneuploidy and polyploidy in Saxifragaceae.     

Molecular evidence for polyploid origins in Saxifraga (Saxifragaceae): the narrow arctic endemic S. svalbardensis and its widespread allies

The recently described polyploid Saxifraga svalbardensis is endemic to the arctic archipelago of Svalbard. We investigated relationships among four closely related species of Saxifraga in Svalbard and tested three previously proposed hypotheses for the origin of S. svalbardensis: (1) differentiation from the morphologically and chromosomally variable polyploid S. cernua; (2) hybridization between the diploid S. hyperborea and S. cernua; and (3) hybridization between the tetraploid S. rivularis and S. cernua. Fifteen populations were analyzed using random amplified polymorphic DNAs (RAPDs) and nucleotide sequences of the chloroplast gene matK and the internal transcribed spacers (ITS) of nuclear ribosomal DNA (rDNA). RAPD and matK data suggest that S. svalbardensis has originated from a hybrid with S. rivularis as the maternal parent and S. cernua as the paternal parent, possibly a single time, whereas ITS data could not be used to discriminate among the hypotheses. The data also suggest that the diploid S. hyperborea is a progenitor of the tetraploid S. rivularis. The four populations examined of S. svalbardensis were virtually identical for RAPD and ITS markers, whereas S. cernua showed high levels of variation, suggesting that the latter polyploid either has formed recurrently or has undergone considerable differentiation since its origin.     

A new endemic species of Saxifraga L. from Spain

A recently erected species of Saxifraga (section Porphyrion ) endemic to the Picos de Europa (Cordillera Cantábrica) is described and illustrated. S. felineri P. Vargas resembles S. aretioides and S. ferdinandi-coburgi , although the habit and leaf shape are the main characters which support its identity. The taxonomic relationships of the three species are discussed, as well as aspects of the distribution and ecology of S. felineri.     

Chemotaxonomic importance of diarylheptanoids and phenylpropanoids in Saxifraga tangutica (Saxifragaceae)

A phytochemical investigation of Saxifraga tangutica led to the isolation of 11 compounds, including eight diarylheptanoids (1–6, 10 and 11) and three phenylpropanoids (7–9). The chemical structures were established by extensive analysis of their MS and NMR spectroscopic data or comparison with literature data. In the present research, we report the isolated compounds 1–11, for the first time, in the species S. tangutica. Moreover, compounds 1, 2 and 4–11 have not been reported from any species in Saxifragaceae family. Furthermore, we discuss the chemotaxonomic significance of the isolated compounds.     

On the gynodioecious polymorphism in Saxifraga granulata L. (Saxifragaceae)

Sexual and vegetative fitness components in hermaphrodite and female plants of the self-compatible, perennial herb Saxifraga granulata are compared using material derived from a gynodioecious population in northern England.
Females produced only 57% as many seeds as hermaphrodites, but their ovule offspring were 1.28 times as fit as those of hermaphrodites, and females were more vegetatively vigorous. The advantages to females in ovule offspring quality and in vegetative reproduction counteract their disadvantages in pollen and seed production and therefore probably play a role in the maintenance of the gynodioecious polymorphism. Pollination ecology, resource reallocation and inbreeding depression all appear to contribute to the observed sex differences in fitness.     

Revisiting of Carex sect. Confertiflorae s.l. (Cyperaceae): New data from molecular and morphological evidence and first insights on Carex biogeography in East Asia

Carex sect. Confertiflorae s.l. is a medium-sized species group (ca. 40 species) with its center of diversity in E Asia (China and Japan). According to morphological traits, the section has been proposed to split into two sections (sects. Confertiflorae sensu Ohwi and Molliculae Ohwi) up to five different ones (sects. Confertiflorae s.s., Molliculae, Dispalatae Ohwi, Ischnostachyae Ohwi, and Alliiformes Akiyama). Recent phylogenetic reconstructions showed Confertiflorae s.l. not to be monophyletic, as species traditionally considered part of it were found to belong to other clades, whereas species traditionally ascribed to other sections were nested within it. In this study, we investigated the phylogenetic structure, morphological affinities, and biogeographic history of sect. Confertiflorae s.l. We employed a taxon-based approach to explore the morphological affinities of the species considered in sect. Confertiflorae and compared the micromorphology of the nutlets of almost all the taxa using SEM. We included 40 samples representing 31 species/subspecies of sect. Confertiflorae s.l. and used two nuclear (ETS and ITS) and three plastid (trnL-F, matK, and rpl32-trnL ^UAG) molecular markers to reconstruct the phylogeny of the group. The phylogenetic analyses confirmed the polyphyly of sect. Confertiflorae s.l., whose representatives were found within five distinct clades. From these, two clades, sect. Confertiflorae and sect. Molliculae, were found to be closely related and contained the majority of the species. The composition of the two clades agreed with the morphological structure of the group, and we confirmed an exclusive combination of features (namely color of basal sheaths, presence of bract sheath, peduncle of lowest spike, inflorescence sex distribution, shape of pistillate glume apex, and color and veins of utricle, among others) characterizing each of the two clades. The origin of the two clades was found to be in the early Pliocene; however, the majority of the diversification events within each clade took place during the Pleistocene. This illustrates that although Asia has been regarded as having little potential ecological space for Carex to diversify due to its climate stability, groups of sedges sub-endemic from that area may have a fairly recent origin related to glaciations. We proposed the rearrangement of sect. Confertiflorae as previously conceived as three independent sections: the monotypic Alliiformes, sect. Molliculae, and sect. Paludosae.     

Haplotype and ribotype diversity of Saxifraga cuneifolia s. l. (Saxifragaceae)

Saxifraga cuneifolia L. (sect. Gymnopera, Saxifragaceae) is a plant distributed in the main mountain ranges of southern Europe, from the eastern Pyrenees to the eastern Carpathian. Currently, two subspecies are recognized based on morphological characteristics: S. cuneifolia subsp. cuneifolia grows in the Maritime Alps and North Apennines and S. cuneifolia subsp. robusta is located in the remaining area of distribution. A more delicate form and a smaller number of flowers in S. cuneifolia subsp. cuneifolia are the morphological characteristics that differentiate this subspecies from the widespread S. cuneifolia subsp. robusta. To explore the genetic diversity and the subspecific geographic patterns of S. cuneifolia s. l. we conducted a molecular study of nuclear and plastid sequences. Samples of S. cuneifolia s. l. have been analysed throughout the distribution area of this species. Our results, based on nuclear (ITS) and plastid (rbcL, trnL–F, and psbA–trnH) markers, showed a genetic characterization of both subspecies presenting discriminant haplotypes and ribotypes that confirm the current subspecific systematics.     

Intercontinental disjunctions between eastern Asia and western North America in vascular plants highlight the biogeographic importance of the Bering land bridge from late Cretaceous to Neogene

This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.     

Phylogenetic analysis of Toxicodendron （Anacardiaceae） and its biogeographic implications on the evolution of north temperate and tropical intercontinental disjunctions

Toxicodendron is a genus in the Rhus complex of Anacardiaceae with a disjunct distribution between eastern Asia and North America, extending to southeastern Asia and the neotropics. Nuclear （internal transcribed spacer, external transcribed spacer, and NIA-i3） and chloroplast （ndhF and trnL-F） sequences were used to construct phylogenetic relationships of Toxicodendron. Phylogenetic analysis of these data strongly support Toxieodendron as a monophyletic group distinct from other genera of the Rhus complex, and the phylogeny does not fully corroborate classification at the sectional level. Two temperate disjunct lineages were detected, one from section Toxicodendron and the other between the eastern North American Toxicodendron vernix and the eastern Asian Toxicodendron vernieifluum. Their divergence times were estimated to be 13.46 （7.95-19.42） and 7.53 （2.76-12.86） mya, respectively. The disjunction between section Griffithii （taxa from warm temperate to tropical Asia） and Toxieodendron striatum （from the neotropics） was supported and their divergence time was estimated to be 20.84 （11.1 6-30.52） mya in the early Miocene. Our biogeographic results and the paleontological data support the Bering land bridge as the most likely route to explain the temperate disjunctions, yet the tropical disjunction in Toxicodendron seems to be best explained by the North Atlantic land bridge hypothesis.     

Phylogeny and biogeography of Aralia sect. Aralia (Araliaceae)

Aralia sect. Aralia (Araliaceae) consists of approximately eight species disjunctly distributed in Asia and North America. Phylogenetic and biogeographic analyses were conducted using sequences of the internal transcribed spacer regions of the nuclear ribosomal DNA. Aralia racemosa from eastern North America was sister to A. californica from western North America. Aralia cordata from eastern Asia did not form a species-pair relationship with the eastern North American A. racemosa. The two subspecies of A. racemosa formed a monophyletic group. Biogeographic analyses showed a close area relationship between eastern North America and western North America. The Himalayas were cladistically basal and eastern Asia was placed between the Himalayas and North America. The biogeographic analysis supported the origin of the eastern Asian and eastern North American disjunct pattern in Aralia sect. Aralia via the Bering land bridges. Comparisons with results of phylogenetic analyses of other genera suggested that (1) the floristic connection between eastern North America and western North America may be stronger than previously thought; and (2) the biogeographic patterns in the Northern Hemisphere are complex. Furthermore, a lack of correlation between sequence divergence values and phylogenetic positions was observed, suggesting the importance of a phylogenetic framework in biogeographic analyses.     

A phylogenetic study of two recently described endemic species of the Saxifraga granulata group from the central-north Mediterranean region (Italy) and their position in the context of the series Saxifraga (Saxifragaceae)

On two small islands of the Tuscan Archipelago, two new endemic species of the Saxifraga granulata group, namely S. caprariae and S. montis-christi, were recently described on a morphological basis. This led us to investigate these species using a molecular approach involving the analysis of ITS and trnL-F regions. Our aims were: to highlight their phylogenetic traits; to confirm or reject the hypothesis that they are more closely related to S. corsica than to S. granulata; and to verify their phylogenetic position in the context of Saxifraga L. ser. Saxifraga. We considered all the species belonging to the ser. Saxifraga and some other species of sect. Saxifraga – a total of 19 species. We generated 10 new DNA sequences, six for ITS and four for trnL-trnF. We produced separate Bayesian and Maximum likelihood trees for both ITS and trnL-trnF regions and we analysed the genetic distances among the new endemics and the more closely related species. The phylogenetic analyses showed that S. caprariae and S. montis-christi are relatively well separated from S. granulata and more related to S. corsica. This supports the taxonomic treatment previously proposed. We showed that the results of our phylogenetic analysis are in agreement with our biogeographic knowledge of the Tuscan Archipelago and the neighbouring territories and emphasize the conservation relevance of these findings.     

基于形态性状的葱属粗根组植物系统发育关系

以环地中海地区和北美葱属Allium的Amerallium亚属种类为外类群,基于36个形态学性状对葱属粗根组sect. Bromatorrhiza进行系统发育分析。简约分析获得的分支拓扑结构与分子系统学的研究结果相吻合,粗根组与环地中海地区的Amerallium亚属种类形成姐妹群关系,表明它们具有密切的亲缘关系。粗根组明显分为两支,粗根韭A. fasciculatum (x = 10,2胚珠)与染色体基数为11的种类(单胚珠)形成一支,另一支由染色体基数为7的大花韭A. macranthum与多星韭A. wallichii (2胚珠)组成,从而否定了粗根亚组subsect. Bromatorrhiza(粗根韭+大花韭+多星韭)的成立。综合已有研究结果,本研究认为粗根组内存在4个相对独立的演化支系,即多星韭及其变种、大花韭、粗根韭,以及染色体基数为11的种类(5种1变种),尽管前人建议的单胚珠亚组subsect. Uniovulatum是合理的,但没有必要在物种数量相对较少的粗根组下进一步细分。     

Evolutionary history of Gymnocarpos (Caryophyllaceae) in the arid regions from North Africa to Central Asia

Gymnocarpos has only about ten species distributed in the arid regions of Asia and Africa, but it exhibits a geographical disjunction between eastern Central Asia and western North Africa and Minor Asia. We sampled eight species of the genus and sequenced two chloroplast regions (rps16 and psbB–psbH), and the nuclear rDNA (ITS) to study the phylogeny and biogeography. The results of the phylogenetic analyses corroborated that Gymnocarpos is monophyletic, in the phylogenetic tree two well supported clades are recognized: clade 1 includes Gymnocarpos sclerocephalus and G. decandrus, mainly the North African group, whereas clade 2 comprises the remaining species, mainly in the Southern Arabian Peninsula. Molecular dating analysis revealed that the divergence age of Gymnocarpos was c. 31.33 Mya near the Eocene and Oligocene transition boundary, the initial diversification within Gymnocarpos dated to c. 6.69 Mya in the late Miocene, and the intraspecific diversification mostly occurred during the Quaternary climate oscillations. Ancestral area reconstruction suggested that the Southern Arabian Peninsula was the ancestral area for Gymnocarpos. Our conclusions revealed that the aridification since mid‐late Miocene significantly affected the diversification of the genus in these areas.     

A comparative study on karyotypes of 28 taxa in Rubus sect. Idaeobatus and sect. Malachobatus (Rosaceae) from China

Rubus is a taxonomically difficult group and cytological data are expected hopefully to gain insight into the relationships of the genus. In this study the chromosome numbers and karyotypes of 18 taxa from sect. Idaeobatus and 10 taxa from sect. Malachobatus were investigated. Among them, the chromosome numbers of 10 taxa and karyotypes of 26 taxa were reported for the first time and mixoploidy was observed new in the genus. The chromosomes are small in size with a length of less than 3 µm and metacentric (about 90%) or submetacentric. All taxa have karyotypes of “1A” except R. cockburnianus, R. innominatus and R. ellipticus var. obcordatus, which have karyotypes of “2A”. No aneuploids were found in all the 28 taxa studied. Plants of sect. Idaeobatus have diploids with 2n=2x=14, except R. idaeopsis (2n=3x=21) and R. parvifolius (A mixoploid of 2n=2x=14 and 2n=4x=28). However, plants of sect. Malachobatus have tetraploids with 2n=4x=28, except for R. buergeri with 2n=8x=56. In addition, conspicuous karyotype differences existed within the 18 taxa belonging to 11 of 7 subsections in sect. Idaeobatus, and the differences between some species within subsections are greater than that between subsections, while uniform karyotypes within subsections and variable karyotypes between subsections were observed in six of 13 subsections in sect. Malachobatus represented by 10 taxa. Systematic values of the cytological data were discussed for some cases when applicable to the two sections.