Evolution of the Eastern Asian and Eastern North American disjunct genus Symplocarpus (Araceae): insights from chloroplast DNA restriction site data

Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases for the eastern Asian and eastern North American disjunct genus Symplocarpus (Araceae). The cpDNA phylogeny reveals a sister group relationship between S. foetidus from eastern North America and S. renifolius from eastern Asia. The cpDNA divergence between the two intercontinental sister species is 0.61%, which suggests an estimated divergence time of 6.1 million years ago during the late Miocene. The Bering land bridge hypothesis is compatible with the estimated time of divergence for the migration of Symplocarpus between eastern Asia and North America. Furthermore, a single origin of the exothermic spadices in Symplocarpus is suggested by the phylogeny. The cpDNA data also provide independent support for the recognition of three species within the genus.     

Molecular phylogeny and biogeography of Astilbe (Saxifragaceae) in Asia and eastern North America

Phylogenetic analyses were conducted for Astilbe (Saxifragaceae), an Asian/eastern North American disjunct genus, using sequences of nuclear ribosomal internal transcribed spacer (ITS) and plastid matK, trnL‐trnF and psbA‐trnH regions. The monophyly of Astilbe is well supported by both ITS and plastid sequences. Topological incongruence was detected between the plastid and the ITS trees, particularly concerning the placement of the single North American species, A. biternata, which may be most probably explained by its origin involving hybridization and/or allopolyploidy with plastid capture. In Astilbe, all species with hermaphroditic flowers constitute a well‐supported clade; dioecious species form a basal grade to the hermaphroditic clade. Astilbe was estimated to have split with Saxifragopsis from western North America at 20.69 Ma (95% HPD: 12.14–30.22 Ma) in the early Miocene. This intercontinental disjunction between Astilbe and Saxifragopsis most likely occurred via the Bering land bridge. The major clade of Astilbe (all species of the genus excluding A. platyphylla) was inferred to have a continental Asian origin. At least three subsequent migrations or dispersals were hypothesized to explain the expansion of Astilbe into North America, Japan and tropical Asian islands. The intercontinental disjunct lineage in Astilbe invokes a hybridization event either in eastern Asia or in North America. This disjunction in Astilbe may be explained by a Beringian migration around 3.54 Ma (95% high posterior density: 1.29–6.18 Ma) in the late Tertiary, although long‐distance dispersal from eastern Asia to North America is also likely. The biogeographical connection between continental Asia, Taiwan, the Philippines and other tropical Asian islands in Astilbe provides evidence for the close floristic affinity between temperate or alpine south‐western China and tropical Asia. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Molecular phylogenetics and biogeography of the eastern Asian–eastern North American disjunct Mitchella and its close relative Damnacanthus (Rubiaceae,Mitchelleae)

Mitchella is a small genus of the Rubiaceae with only two species. It is the only herbaceous semishrub of the family showing a disjunct distribution in eastern Asia and eastern North America, extending to Central America. Its phylogeny and biogeographical diversification remain poorly understood. In this study, we conducted phylogenetic and biogeographical analyses for Mitchella and its close relative Damnacanthus based on sequences of the nuclear internal transcribed spacer (ITS) and four plastid markers (rbcL, atpB‐rbcL, rps16 and trnL‐F). Mitchella is monophyletic, consisting of an eastern Asian M. undulata clade and a New World M. repens clade. Our results also support Michella as the closest relative to the eastern Asian Damnacanthus. The divergence time between the two intercontinental disjunct Mitchella species was dated to 7.73 Mya, with a 95% highest posterior density (HPD) of 3.14?12.53 Mya, using the Bayesian relaxed clock estimation. Ancestral area reconstructions suggest that the genus originated in eastern Asia. The semishrub Mitchella appears to have arisen from its woody ancestor in eastern Asia and then migrated to North America via the Bering land bridge in the late Miocene. © 2013 The Linnean Society of London     

Intercontinental biogeography of subfamily Orontioideae (Symplocarpus, Lysichiton, and Orontium) of Araceae in Eastern Asia and North America

Symplocarpus, Lysichiton, and Orontium (Orontioideae) are three of the few north temperate genera of the primarily tropical Araceae. Symplocarpus is disjunctly distributed in eastern Asia (3 spp.) and eastern North America (1 sp.); Lysichiton has an intercontinental discontinuous distribution in eastern Asia (1 sp.) and northwestern North America (1 sp.); and the monotypic Orontium is restricted to eastern North America. Phylogenetic analysis of the trnL-F and ndhF sequences supports (1) the monophyly of both Symplocarpus and Lysichiton, (2) the sister-group relationship of Symplocarpus and Lysichiton, and (3) the clade of Orontium, Symplocarpus, and Lysichiton. Although Symplocarpus shows a much wider disjunction than Lysichiton, the estimated divergence time of the former [4.49+/-1.69 or 6.88+/-4.18 million years ago (mya)] was similar to that of the latter (4.02+/-1.60 or 7.18+/-4.33 mya) based on the penalized likelihood and the Bayesian dating methods, respectively. Eastern Asia was suggested to be the ancestral area of the Symplocarpus-Lysichiton clade based on the dispersal-vicariance analysis. Our biogeographic results support independent migrations of Symplocarpus and Lysichiton across the Bering land bridge in the late Tertiary (Pliocene/late Miocene). Fossil evidence suggests Orontioideae dated back to the late Cretaceous in the temperate Northern Hemisphere (72 mya). The relative rate test shows similar substitution rates of the trnL-F sequences between the proto and the true aroids, although the latter has substantially higher species diversity. The proto Araceae perhaps suffered from a higher rate of extinction in the temperate zone associated with periods of climatic cooling in the Tertiary.     

New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history

The heat‐ and odour‐producing genus Arum (Araceae) has interested scientists for centuries. This long‐term interest has allowed a deep knowledge of some complex processes, such as the physiology and dynamics of its characteristic lure‐and‐trap pollination system, to be built up. However, mainly because of its large distributional range and high degree of morphological variation, species' limits and relationships are still under discussion. Today, the genus comprises 28 species subdivided into two subgenera, two sections and six subsections. In this study, the phylogeny of the genus is inferred on the basis of four plastid regions, and the evolution of several morphological characters is investigated. Our phylogenetic hypothesis is not in agreement with the current infrageneric classification of the genus and challenges the monophyly of several species. This demonstrates the need for a new infrageneric classification based on characters reflecting the evolution of this enigmatic genus. To investigate the biogeography of Arum deeply, further spatiotemporal analyses were performed, addressing the importance of the Mediterranean basin in the diversification of Arum. Our results suggest that its centre of origin was the European–Aegean region, and that major diversification happened during the last 10 Myr. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 14–32.     

Size-dependent flowering behavior and heat production of a sequential hermaphrodite, Symplocarpus renifolius (Araceae)

We examined the flowering performance in a population of the protogynous perennial herb Symplocarpus renifolius (Araceae), with special consideration of plant size. Flowering of S. renifolius occurred in very early spring, soon after snow melt. The spadices generated heat throughout the pistillate (female) and bisexual phases, but heat production decreased quickly after the beginning of the staminate (male) phase. During the flowering season, the sex ratio within the population dramatically changed from a dominance of females to a dominance of males. The duration of the female phase was negatively correlated with the onset time of flowering, and the duration of the male phase was positively correlated with plant size. Larger plants began blooming earlier, produced more heat, made the transition from female to male phase more rapidly, and lasted longer as males than smaller ones. Such size-dependent flowering performance caused unidirectional pollen flow from large to small plants. The number of seeds produced per spadix was positively correlated with the duration of the female phase, although it was not correlated with plant size. However, the estimated number of seeds sired during the male phase was positively correlated with plant size. Early flowering, rapid gender change, and higher heat production of the spadices by larger plants were factors considered to promote the higher success of the male function without decreasing the success of the female function.     

Cytology and systematics in JapaneseArisaema (Araceae)

Chromosome numbers are determined from 37 populations attributed to 22 taxa of JapaneseArisaema. Of them, chromosome numbers ofA. limbatum var.conspicuum (2n=26),A. minus (2n=26),A. nambae (2n=28) andA. seppikoense (2n=26) are determined for the first time. New chromosome numbers, 2n=26, are reported forA. aequinoctiale, A. limbatum, A. stenophyllum, A. undulatifolium andA. yoshinagae. Three modes of basic chromosome numbers,x=14,x=13 andx=12, occur in JapaneseArisaema. Precise karyotypic comparisons of 20 taxa reveal that taxa withx=14 andx=13 share 26 major chromosome arms and have an obvious chromosomal relationship. One of two submeta-centric chromosomes inx=13 corresponds to two telo-centric chromosomes inx=14. InA. ternatipartitum with 2n=6x=72, ten out of 12 basic chromosomes are the most similar in size and arm ratio with larger ten chromosomes ofA. ringens among JapaneseArisaema examined. A basic chromosome number ofx=14 is the commonest in the genusArisaema and the remaining basic chromosome numbers,x=13 andx=12, seem to be derived through dysploidal reduction by translocating large segments of major arm of telo-centric chromosome onto other minor arm of telo-centric followed by loss of the remainings including a centromere, and by loss of two telo-centrics fromx=14, respectively. Some systematic problems of JapaneseArisaema are discussed based on new cytological data.Arisaema hatizyoense, A. minus andA. nambae are accepted as independent species.     

六种犁头尖属植物(天南星科)的核型研究

报道了 6种 8个居群犁头尖属 ( Typhonium Schott)植物的核型 ,其结果如下 :( 1 )独角莲 ( T.gigan-teum)北京居群 2 n=4 x=5 2 =4 4m+ 7sm+ 1 st;( 2 )鞭檐犁头尖 ( T.flagelliforme)金平居群 2 n=3x=2 4 =3m+ 9sm( 4 SAT) + 1 2 st,河内居群 2 n=4 x=32 =7m+ 2 0 st+ 4sm+ 1 t;( 3)单籽犁头尖 ( T. calcicolum)西畴居群2 n=4 x=5 2 =2 1 sm+ 2 3m( 5 SAT) + 8st;( 4 )犁头尖 ( T.blumei)重庆居群 2 n=4 x=5 2 =4 0 m( 1 SAT) + 1 2 sm( 3SAT) ;( 5 )马蹄犁头尖 ( T.trilobatum)西双版纳居群 2 n=2 x=1 8=4 sm( 2 SAT) + 1 2 m+ 2 st,河内居群 2 n=2 x=1 8=2 st+ 9m+ 7sm;( 6 )金慈菇 ( T. roxburgii)个旧居群 2 n=2 x=1 8=8sm+ 1 0 m( 2 SAT)。其中鞭檐犁头尖 2 n=2 4、32 ,金慈菇 2 n=1 8均为首次报道 ,同时分析讨论了本属植物染色体基数和倍性的多样性及其可能的原始基数     

Flavonoids and xanthones from the leaves of Amorphophallus titanum (Araceae)

The flavonoids and xanthones in the leaves of Amorphophallus titanum, which has the largest inflorescence among all Araceous species, were surveyed. Eight C-glycosylflavones, five flavonols, one flavone O-glycoside and two xanthones were isolated and characterized as vitexin, isovitexin, orientin, isoorientin, schaftoside, isoschaftoside, vicenin-2 and lucenin-2 (C-glycosylflavones), kaempferol 3-O-robinobioside, 3-O-rutinoside and 3-O-rhamnosylarabinoside, and quercetin 3-O-robinobioside and 3-O-rutinoside (flavonols), luteolin 7-O-glucoside (flavone), and mangiferin and isomangiferin (xanthones). Although the inflorescence of this species has been surveyed for flavonoids, those of the leaves were reported for the first time.     

国产芋属花粉形态

通过光镜和扫描电子显微镜观察了芋属(Colocasia Schott)6种、1变种、1品种的花粉形态,其中贡山芋C．gaoligongensis,龚氏芋C．gongii,李氏香芋C．lihengiae,花叶芋C．bic-olor4个种为首次报道。本属花粉为圆球形,无萌发孔。外壁纹饰分为两类：大野芋C．giga-ntea外壁光滑无刺,其余各种均具刺。花粉大小为20．07—32．76μm。     

Molecular phylogeny and historical biogeography of Caltha (Ranunculaceae) based on analyses of multiple nuclear and plastid sequences

Caltha is a widely distributed genus in the buttercup family (Ranunculaceae) showing interesting distribution patterns in both hemispheres. Evolutionary history ofCaltha was examined by means of phylogenetic, molecular dating, and historical biogeographic analyses with a more comprehensive sampling than previous studies. The internal transcribed spacer from the nuclear genome and trnL-F and atpB-rbcL regions from the plastid genome were used and analyzed using parsimony and Bayesian methods. Divergence time was estimated using Bayesian dating analyses with multiple fossil calibrations. Historical biogeography was inferred using the Bayes-DIVA method implemented in RASP. We obtained a well-resolved and well-supported phylogeny within the Caltha lineage. Caltha natansPall. diverged first from the genus and the other species grouped into two clades. Our expanded sampling scheme revealed a complicated evolutionary pattern in theC. palustris complex. Caltha sinogracilis W. T. Wang was resolved to be a member of the C. palustris complex, rather than closely related to C. scaposa Hook. f. & Thomson. Caltha rubriflora B. L. Burtt & Lauener was also revealed to be not just a red-flower form of C. sinogracilis. The diversification of the genus began at 50.5 mya (95% high posterior density: 37.1–63.9 mya), and its ancestral range was very probably in the Northern Hemisphere. The South American species may derive from western North American ancestors that dispersed along the western American Cordillera during the Cenozoic era. The vicariance model of the Southern Hemisphere species proposed by a previous study was rejected in this study.     

Araceae of Mulu National Park. I. Four new species of Schismatoglottis (Araceae)

Four new species of Schismatoglottis are described and illustrated from Mulu National Park in Malaysian Borneo: Schismatoglottis fossae (Patentinervia Clade), S. roseopedes and S. serratodentata (Calyptrata Clade) and S. pellucida (currently unplaced but likely in the Multinervia clade). A brief summary is provided of botanical exploration at Mulu, notably where pertinent to aroid botany.     

Maximum likelihood inference implies a high, not a low, ancestral haploid chromosome number in Araceae, with a critique of the bias introduced by 'x'

Background and Aims

For 84 years, botanists have relied on calculating the highest common factor for series of haploid chromosome numbers to arrive at a so-called basic number, x. This was done without consistent (reproducible) reference to species relationships and frequencies of different numbers in a clade. Likelihood models that treat polyploidy, chromosome fusion and fission as events with particular probabilities now allow reconstruction of ancestral chromosome numbers in an explicit framework. We have used a modelling approach to reconstruct chromosome number change in the large monocot family Araceae and to test earlier hypotheses about basic numbers in the family.

Methods

Using a maximum likelihood approach and chromosome counts for 26 % of the 3300 species of Araceae and representative numbers for each of the other 13 families of Alismatales, polyploidization events and single chromosome changes were inferred on a genus-level phylogenetic tree for 113 of the 117 genera of Araceae.

Key Results

The previously inferred basic numbers x = 14 and x = 7 are rejected. Instead, maximum likelihood optimization revealed an ancestral haploid chromosome number of n = 16, Bayesian inference of n = 18. Chromosome fusion (loss) is the predominant inferred event, whereas polyploidization events occurred less frequently and mainly towards the tips of the tree.

Conclusions

The bias towards low basic numbers (x) introduced by the algebraic approach to inferring chromosome number changes, prevalent among botanists, may have contributed to an unrealistic picture of ancestral chromosome numbers in many plant clades. The availability of robust quantitative methods for reconstructing ancestral chromosome numbers on molecular phylogenetic trees (with or without branch length information), with confidence statistics, makes the calculation of x an obsolete approach, at least when applied to large clades.     

Reasons and consequences of the lack of a sporopollenin ektexine in Aroideae (Araceae)

The ultrastructure of pollen walls in Araceae is characterized by the absence of a stable sporopollenin outer exine layer in subfamily Aroideae, and by the presence of several distinctive pollen characters typical for the other aroid subfamilies. This article discusses if and to which extent such distinctive pollen characters are mirrored in various classifications of Araceae, basing either on morphological or on molecular data. Accordingly, the pollen characters perfectly reflect the actual subfamily classification, and also recent arrangements of clades in trees basing on molecular data. The actual subfamilies appear no longer eurypalynous, but now strictly stenopalynous. Aside from the (settled) classification problem the fundamental question is addressed why do Aroideae lack an elaborated sporopollenin ektexine. Possible pollination biology benefits, deriving from an absence of an elaborated sporopollenin ektexine in Aroideae, are presented and discussed. Compared with all other subfamilies the most advanced and by far largest subfamily Aroideae has lost several crucial characters and simultaneously acquired corresponding opposed characters, amongst others a non-sporopollenin exine layer and an unusual thick and spongy endexine. Taken together, losses and acquisitions are interpreted as a major paradigm shift in Araceae evolution, which took place according to the fossil record probably in the Paleogene.     

斑龙芋属(天南星科)及近缘属植物的叶表皮形态

光镜、电镜下观察了天南星科天南星族下斑龙芋属2个种及近缘属13个种植物的叶表皮形态特征。实验结果显示,4属植物的叶表皮组成及其形态特征较相似,属间不存在明显差异,但某些特征在种间存在差异,可作为种的鉴别特征。叶表皮特征支持将单籽犁头尖和昆明犁头尖两个种合并为一个种。15个种的气孔器均具有2个副卫细胞,Stebbins andKhush认为这是气孔器类型中较具2个以上副卫细胞更进化的一种类型,而天南星科大多数族的气孔器都具有2个以上的副卫细胞,这也证明了天南星族是天南星科较进化的族。     

The uneven phylogeny and biogeography of Erodium (Geraniaceae): radiations in the Mediterranean and recent recurrent intercontinental colonization

Background and Aims

The genus Erodium is a common feature of Mediterranean-type climates throughout the world, but the Mediterranean Basin has significantly higher diversity than other areas. The aim here is to reveal the biogeographical history of the genus and the causes behind the evolution of the uneven distribution.

Methods

Seventy-eight new nrITS sequences were incorporated with existing plastid data to explore the phylogenetic relationships and biogeography of Erodium using several reconstruction methods. Divergence times for major clades were calculated and contrasted with other previously published information. Furthermore, topological and temporal diversification rate shift analyses were employed using these data.

Key Results

Phylogenetic relationships among species are widely congruent with previous plastid reconstructions, which refute the classical taxonomical classification. Biogeographical reconstructions point to Asia as the ancestral area of Erodium, arising approx. 18 MYA. Four incidences of intercontinental dispersal from the Mediterranean Basin to similar climates are demonstrated. Increases in diversification were present in two independent Erodium lineages concurrently. Two bursts of diversification (3 MYA and 0·69 MYA) were detected only in the Mediterranean flora.

Conclusions

Two lineages diverged early in the evolution of the genus Erodium: (1) subgenus Erodium plus subgenus Barbata subsection Absinthioidea and (2) the remainder of subgenus Barbata. Dispersal across major water bodies, although uncommon, has had a major influence on the distribution of this genus and is likely to have played as significant role as in other, more easily dispersed, genera. Establishment of Mediterranean climates has facilitated the spread of the genus and been crucial in its diversification. Two, independent, rapid radiations in response to the onset of drought and glacial climate change indicate putative adaptive radiations in the genus.     

Upper Cretaceous sulcate pollen from the Timerdyakh Formation,Vilui Basin (Siberia)

One inaperturate and 16 monosulcate pollen types are described from the latest Campanian to earliest Maastrichtian sediments of the Vilui basin, Siberia, using both light and scanning electron microscopy, and assigned systematically when possible to modern families or subfamilies. Despite their scant occurrence, the pollen grains show considerable diversity. Two new genera have been erected. Lasioideaecidites with two species, Lasioideaecidites hessei and Lasioideaecidites bogneri, represents the earliest record of the subfamily Lasioideae (Araceae). Aristolochiacidites with one species, Aristolochiacidites viluiensis, is assigned to the subfamily Aristolochioideae (Aristolochiaceae) and represents the first fossil pollen record of the family. A new species of Liliacidites, Liliacidites goldblattii, is closely similar to pollen of Isophysioideae (Iridaceae) and/or Doryanthaceae and is the first fossil evidence of the clade that includes these closely related families. A further nine new fossil species are described (Clavatipollenites timerdyakhensis, Liliacidites palaeofritillaria, Retimonocolpites longosucatus, R. microreticulatus, R. microrugulatus, R. lysichitonoides, R. chapmaniae, Monosulcites parvus and Arecipites tyungensis) and assigned to Chloranthaceae, Araceae (Orontioideae, ?Pothoideae), Liliaceae, ?Hypoxidaceae and Arecaceae. The Hydatellaceae (Nymphaeales) may also be present in the palynoflora of the Vilui Basin, represented by pollen similar to M. rivularis Braman from the Santonian to Campanian (Upper Cretaceous) Milk River Formation, Canada.     

Chemical constituents from the tubers of Pinellia ternata (Araceae) and their chemotaxonomic interest

This work describes the isolation and characterization of twenty compounds from the tubers of Pinellia ternata (Thunb.) Breit, including 17 lignanoids (1–17) and three alkaloids (18–20). All structures were established based on analysis of their spectroscopic data. It is the first report of compounds 3–11 and 13–18 from the genus Pinellia, and the first report of compounds 3–4, 6–9, 11 and 13–18 from the family Araceae. In addition, the biosynthetic pathways of the 17 lignanoids are summarized. A liquid chromatography coupled with electrospray ionization quadrupole time-of-flight mass spectrometry (LC-ESI-QTOFMS/MS) method in both positive and negative ion mode was also used to confirm the identity of these seventeen lignanoids from the tubers of P. ternata. The results from the present work provide further information about the diversity of compounds in the genus Pinellia.