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Two new species of Impatiens (Balsaminaceae) from Gaoligong Mountains, Yunnan, China
Abstract
Impatiens longyangensis and I. yaojiapingensis, two new species from Gaoligong Mountains, Yunnan, China, are described here. The former species is morphologically similar to I. shangjiangensis, but differs by its shorter petioles and peduncle, ovate bract, shortly saccate lower sepal with a longer spur and suborbicular dorsal petal. The latter is similar to I. sterilis, and differs by its pubescent leaf, petiole with one to three pairs of clavate glands, persistent bract, lower sepal with a straight spur and suborbicular dorsal petal. Molecular phylogenetic analysis based on nuclear ITS and plastid atpB-rbcL sequences shows that the two new species belong to I. sect. Racemosae.
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Impatiens nushanensis (Balsaminaceae), a new species from Yunnan, China, is described and illustrated here. The new species is morphologically most similar to I. purpurea, but differs by having two pairs of lateral sepals and deep purple lower lobe of the lateral united petal with an acuminate and ribbon-like apex. Furthermore, phylogenetic analysis based on molecular data also support the recognization of this new species.
Impatiens longiaristata (Balsaminaceae), a new species from western Sichuan Province in China, is described and illustrated here based on morphological and molecular data. It is similar to I. longiloba and I. siculifer, but differs in its lower sepal with a long arista at the apex of the mouth, spur curved downward or circinate, and lower petal that is oblong-elliptic and two times longer than the upper petal. Molecular analysis confirmed its placement in sect. Racemosae. Simultaneously, during the inspection of the protologues and type specimens of allied species, it was found that the types of two names from this section were syntypes based on Article 9.6 of the International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code). According to Articles 8.1, 9.3, and Recommendations 9A.1, 9A.2, and 9A.3, the lectotypes of these two names are here designated.
Impatiens longshanensis (The LSID for the name Impatiens longshanensis is: 77219154-1) sp. nov. and I. lihengiana (The LSID for the name I. lihengiana is: 77219153-1) sp. nov., from Hunan, China, are described and illustrated here. The molecular phylogenetic study suggests that I. longshanensis and I. lihengiana should be placed in the I. sect. Impatiens. A detailed description, diagnostic characters between the two new species and allied species, pollen and seed morphology, and color photographs are provided. In addition, based on wide sampling, we found that the longifilamenta group, an endemic group to China, whose members have basal lobes of lateral united petals with long filamentous hairs, shows significant morphological variability. In this paper, we discuss the taxonomic significance of morphological characteristics within this group. Based on a literature review and observation of living materials in the field, an updated identification key for this group is also proposed.
Impatiens rapiformis (Balsaminaceae), a new species from Malipo County, Yunnan, China, is described and illustrated. The new species is superficially similar to I. apalophylla, I. malipoensis and I. guizhouensis, but differs by having cylindric or subfusiform root tuber, elliptic or obovate‐oblong leaf blades and falcate outer lateral sepals. The characters of its seed micromorphology and pollen morphology under SEM are given. A preliminary molecular phylogenetic study suggests that I. rapiformis should be placed in I. subg. Clavicarpa.
Impatiens bullatisepala (Balsaminaceae), a new species supported by morphological and phylogenetic evidence from Fanjing Mountain, Guizhou province in China, is described here. It is morphologically similar to I. davidii but can be distinguished by its dorsally ridged lateral sepals with sunk reticulate veins and bullate projections on abaxial surface, 2–2.5 cm deep saccate lower sepal with ca. 0.8 cm long narrowly triangular tip at the mouth, and broadly ovate dorsal petal. Phylogenetic analyses of a combined dataset of nuclear ITS and plastid atpB-rbcL intergenic spacer DNA sequences furtherly confirmed its novelty.
Impatiens wutaishanensis R.L. Liao & Lei Cai, a new species from Southeast Yunnan, China, is here described and illustrated. This new species is most similar to Impatiens parvisepala S.X. Yu & Y.T. Hou in its racemose inflorescences, its four lateral sepals, the leaf arrangement, and in having yellow flowers. However, it differs in the height of the plants, the length of the petiole, the size and shape of the leaf blade, the shape of the spur, and the number of flowers in each inflorescence. A detailed description, color photographs, and a provisional IUCN red list assessment are provided, and its geographical distribution, ecology, and morphological relationship with relevant similar species are discussed.
Background
Southeast Asia, together with tropical Africa, Madagascar, South India and Sri Lanka, and the eastern Himalayas, are the five primary hotspots of species diversity of Impatiens (Balsaminaceae). China is also rich in Impatiens species, especially in the limestone karsts or ‘Danxia’ landforms. With zygomorphic flowers and diverse corolla morphology and color, the species in Impatiens are well-known for their ornamental use, but they are also notorious in taxonomy. During the preparation of revision of Impatiens in Zhejiang and adjacent regions, an unknown species was collected from Mt. Wuyi in Fujian Province, Southeast China.
Results
Phylogenetic analyses based on nuclear ITS, chloroplast atp B- rbc L and trn L-F sequences, together with micromorphology of pollen grains and seed coats, strongly supported the close relationship of the new species with Impatiens platysepala Y.L.Chen and I. chloroxantha Y.L.Chen. In turn, both molecular data and morphological characters also were sufficient to distinguish the new species from the other two counterparts.
Conclusions
Our detailed morphological observations and molecular phylogenetic analyses support the recognition of Impatiens wuyiensis as a species new to science.
Impatiens quintadecimacopii G. W. Hu & Q. F. Wang sp. nov. is described and I. porrecta is reported as a new record from southwestern Yunnan province of China. These two species are close in morphology but can be distinguished by color of flowers, shape of lateral sepals and upper petals, and color of capsules. Morphological characters of these two species indicate they belong to sect. Uniflora in I. subg. Impatiens. Phylogenetic analysis of a combined dataset from nuclear ITS and plastid atpB-rbcL sequences confirmed their close relationship and their phylogenetic placement in sect. Uniflora.
Yunnan, located in southwestern China, harbors more than 19,000 higher plants, which represents the highest plant diversity in the country. However, plant diversity in Yunnan faces enormous threats today, including habitat destruction and fragmentation, environmental pollution, and over-exploitation of natural resources. Despite recent efforts to protect biodiversity, there are still thousands of threatened species, some of which have become extinct. We analyzed available data to gain a greater understanding of plant diversity and the status of plant conservation in Yunnan. We found that southern, southeastern, and northwestern Yunnan are hotspots of total species, endemic species, specimens, new species and threatened species, whereas southeastern Yunnan are hotspots for plant species with extremely small populations. Moreover, we found that there are still conservation gaps and poorly protected areas in central, eastern, and northeastern Yunnan. We conclude that continued modern investigation is required, as are systematics research, the development of comprehensive databases, and government support. We recommend that conservationists pay more attention to building and improving functional protection systems and popularizing science.
Multigene and genomic data sets have become commonplace in the field of phylogenetics, but many existing tools are not designed for such data sets, which often makes the analysis time‐consuming and tedious. Here, we present PhyloSuite, a (crossplatform, open‐source, stand‐alone Python graphical user interface) user‐friendly workflow desktop platform dedicated to streamlining molecular sequence data management and evolutionary phylogenetics studies. It uses a plugin‐based system that integrates several phylogenetic and bioinformatic tools, thereby streamlining the entire procedure, from data acquisition to phylogenetic tree annotation (in combination with iTOL). It has the following features: (a) point‐and‐click and drag‐and‐drop graphical user interface; (b) a workplace to manage and organize molecular sequence data and results of analyses; (c) GenBank entry extraction and comparative statistics; and (d) a phylogenetic workflow with batch processing capability, comprising sequence alignment (mafft and macse), alignment optimization (trimAl, HmmCleaner and Gblocks), data set concatenation, best partitioning scheme and best evolutionary model selection (PartitionFinder and modelfinder), and phylogenetic inference (MrBayes and iq‐tree). PhyloSuite is designed for both beginners and experienced researchers, allowing the former to quick‐start their way into phylogenetic analysis, and the latter to conduct, store and manage their work in a streamlined way, and spend more time investigating scientific questions instead of wasting it on transferring files from one software program to another.
The molecular evolutionary genetics analysis (Mega) software implements many analytical methods and tools for phylogenomics and phylomedicine. Here, we report a transformation of Mega to enable cross-platform use on Microsoft Windows and Linux operating systems. Mega X does not require virtualization or emulation software and provides a uniform user experience across platforms. Mega X has additionally been upgraded to use multiple computing cores for many molecular evolutionary analyses. Mega X is available in two interfaces (graphical and command line) and can be downloaded from www.megasoftware.net free of charge.
Impatiens L. is one of the largest angiosperm genera, containing over 1000 species, and is notorious for its taxonomic difficulty. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of the genus to date based on a total evidence approach. Forty-six morphological characters, mainly obtained from our own investigations, are combined with sequence data from three genetic regions, including nuclear ribosomal ITS and plastid atpB-rbcL and trnL-F. We include 150 Impatiens species representing all clades recovered by previous phylogenetic analyses as well as three outgroups. Maximum-parsimony and Bayesian inference methods were used to infer phylogenetic relationships. Our analyses concur with previous studies, but in most cases provide stronger support. Impatiens splits into two major clades. For the first time, we report that species with three-colpate pollen and four carpels form a monophyletic group (clade I). Within clade II, seven well-supported subclades are recognized. Within this phylogenetic framework, character evolution is reconstructed, and diagnostic morphological characters for different clades and subclades are identified and discussed. Based on both morphological and molecular evidence, a new classification outline is presented, in which Impatiens is divided into two subgenera, subgen. Clavicarpa and subgen. Impatiens; the latter is further subdivided into seven sections.
Large phylogenomics data sets require fast tree inference methods, especially for maximum-likelihood (ML) phylogenies. Fast
programs exist, but due to inherent heuristics to find optimal trees, it is not clear whether the best tree is found. Thus,
there is need for additional approaches that employ different search strategies to find ML trees and that are at the same
time as fast as currently available ML programs. We show that a combination of hill-climbing approaches and a stochastic perturbation
method can be time-efficiently implemented. If we allow the same CPU time as RAxML and PhyML, then our software IQ-TREE found
higher likelihoods between 62.2% and 87.1% of the studied alignments, thus efficiently exploring the tree-space. If we use
the IQ-TREE stopping rule, RAxML and PhyML are faster in 75.7% and 47.1% of the DNA alignments and 42.2% and 100% of the protein
alignments, respectively. However, the range of obtaining higher likelihoods with IQ-TREE improves to 73.3–97.1%. IQ-TREE
is freely available at http://www.cibiv.at/software/iqtree.
The genus Impatiens (Balsaminaceae) is one of the most difficult genera for making effective herbarium specimens because of its tender and complicated flowers. The preparation of flowers is always one of the most important steps in the collections of Impatiens. The present paper demonstrates a method to prepare flowering specimens of Impatiens in the field. Examples include three new species collected from the Sino-Vietnamese border, viz. Impatiens aconitoides, I. purpureifolia, and I. rugata. The related species I. laojunshanensis, I. apalophylla, and I. clavigera are also sampled using the same preparation method. As to the three new species, their pollen grains and seeds were observed by scanning electron microscopy. Impatiens aconitoides has 4-colpate pollen grains and ovoid seeds with protrusive-type of seed coat, whereas I. purpureifolia and I. rugata have 3-colpate grains and ellipsoidal seeds with reticulate-typed seed coat. Such an integrated approach to include detailed floral characters as well as pollen and seed characters is proposed when new taxa of Impatiens are described.
Sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA were acquired for 112 species of Balsaminaceae worldwide and five species of its closest relatives Marcgraviaceae and Tetrameristaceae. Phylogenetic analyses applying parsimony and distance estimates confirmed the monophyly of Balsaminaceae and suggest the monophyly of Impatiens. Within Impatiens, a few clades are recognized with strong support. Two of the most important clades are the spurless Madagascan endemic group, and the one comprising species with broadly fusiform fruits and the basic chromosome number x = 8, that shows a Southeast Asia, southern India, Africa, and Madagascar connection. Despite recognition of several strongly supported small lineages, ITS data alone could not resolve relationships among most of the lineages with confident support values. ITS phylogenies are therefore of limited taxonomic value for Impatiens. However, ITS phylogenies do reveal that extant Impatiens species are of Southeast Asian origin, from where dispersals to boreal Eurasia and North America, to central Asia and eastern Europe via the Himalayas, and to India and Africa have occurred. The Madagascan Impatiens show an African origin. Molecular phylogenies suggest the ancestral basic chromosome number to be x = 10, and the spurred flowers and elongated linear fruits to be plesiomorphic states in Impatiens. A predominantly descending dysploid chromosome evolution, following dispersal of the clade with broadly fusiform fruits from Southeast Asia to India, Africa, and Madagascar, is also suggested.
Balsaminaceae are a morphologically diverse family with ca. 1,000 representatives that are mainly distributed in the Old World tropics and subtropics. To understand the relationships of its members, we obtained chloroplast atpB-rbcL sequences from 86 species of Balsaminaceae and five outgroups. Phylogenetic reconstructions using parsimony and Bayesian approaches provide a well-resolved phylogeny in which the sister group relationship between Impatiens and Hydrocera is confirmed. The overall topology of Impatiens is strongly supported and is geographically structured. Impatiens likely originated in South China from which it colonized the adjacent regions and afterwards dispersed into North America, Africa, India, the Southeast Asian peninsula, and the Himalayan region.
Since its introduction in 2001, MrBayes has grown in popularity as a software package for Bayesian phylogenetic inference using Markov chain Monte Carlo (MCMC) methods. With this note, we announce the release of version 3.2, a major upgrade to the latest official release presented in 2003. The new version provides convergence diagnostics and allows multiple analyses to be run in parallel with convergence progress monitored on the fly. The introduction of new proposals and automatic optimization of tuning parameters has improved convergence for many problems. The new version also sports significantly faster likelihood calculations through streaming single-instruction-multiple-data extensions (SSE) and support of the BEAGLE library, allowing likelihood calculations to be delegated to graphics processing units (GPUs) on compatible hardware. Speedup factors range from around 2 with SSE code to more than 50 with BEAGLE for codon problems. Checkpointing across all models allows long runs to be completed even when an analysis is prematurely terminated. New models include relaxed clocks, dating, model averaging across time-reversible substitution models, and support for hard, negative, and partial (backbone) tree constraints. Inference of species trees from gene trees is supported by full incorporation of the Bayesian estimation of species trees (BEST) algorithms. Marginal model likelihoods for Bayes factor tests can be estimated accurately across the entire model space using the stepping stone method. The new version provides more output options than previously, including samples of ancestral states, site rates, site d(N)/d(S) rations, branch rates, and node dates. A wide range of statistics on tree parameters can also be output for visualization in FigTree and compatible software.
PhyML is a phylogeny software based on the maximum-likelihood principle. Early PhyML versions used a fast algorithm performing nearest neighbor interchanges to improve a reasonable starting tree topology. Since the original publication (Guindon S., Gascuel O. 2003. A simple, fast and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst. Biol. 52:696-704), PhyML has been widely used (>2500 citations in ISI Web of Science) because of its simplicity and a fair compromise between accuracy and speed. In the meantime, research around PhyML has continued, and this article describes the new algorithms and methods implemented in the program. First, we introduce a new algorithm to search the tree space with user-defined intensity using subtree pruning and regrafting topological moves. The parsimony criterion is used here to filter out the least promising topology modifications with respect to the likelihood function. The analysis of a large collection of real nucleotide and amino acid data sets of various sizes demonstrates the good performance of this method. Second, we describe a new test to assess the support of the data for internal branches of a phylogeny. This approach extends the recently proposed approximate likelihood-ratio test and relies on a nonparametric, Shimodaira-Hasegawa-like procedure. A detailed analysis of real alignments sheds light on the links between this new approach and the more classical nonparametric bootstrap method. Overall, our tests show that the last version (3.0) of PhyML is fast, accurate, stable, and ready to use. A Web server and binary files are available from http://www.atgc-montpellier.fr/phyml/.
Impatiens shangjiangensis Y.Y. Cong & J.Z. Gu, a new species of Balsaminaceae from Lushui County, Yunnan, China, is described and illustrated. It is similar to I. bahanensis, but differs from it in having bigger flowers, a curved-saccate lower sepal, and a reniform dorsal petal. Pollen grains are also different in the two species.
Impatiens damingensis S. X. Yu, C. Y. Xia & H. P. Deng, (Balsaminaceae) discovered in Guangxi, China, is described and illustrated here. This species is similar to I. aquatilis Hook.f., but differs from the latter by having blades that are ovate or ovate-lanceolate vs. lanceolate, ovate-lanceolate, or ovate-oblong; apices of bracts acute vs. glandular awned; apices of lateral sepals mucronulate vs. long mucronulate; and spur slender substraight or slightly incurved vs. incurved. Furthermore, molecular data as well as micro-morphological evidence under SEM (of pollens and seeds) also support the establishment of the new species.
Model-based molecular phylogenetics plays an important role in comparisons of genomic data, and model selection is a key step in all such analyses. We present ModelFinder, a fast model-selection method that greatly improves the accuracy of phylogenetic estimates by incorporating a model of rate heterogeneity across sites not previously considered in this context and by allowing concurrent searches of model space and tree space.
Conservationists are far from able to assist all species under threat, if only for lack of funding. This places a premium on priorities: how can we support the most species at the least cost? One way is to identify 'biodiversity hotspots' where exceptional concentrations of endemic species are undergoing exceptional loss of habitat. As many as 44% of all species of vascular plants and 35% of all species in four vertebrate groups are confined to 25 hotspots comprising only 1.4% of the land surface of the Earth. This opens the way for a 'silver bullet' strategy on the part of conservation planners, focusing on these hotspots in proportion to their share of the world's species at risk.
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