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Abstract

The generic boundary of the broadly defined Acanthophyllum s.l., the third‐largest genus of the tribe Caryophylleae (Caryophyllaceae), has been a subject of taxonomic confusion. Acanthophyllum s.l. now includes five minor genera previously recognized as independent. Among these small genera, the inclusion of Allochrusa, Ochotonophila, and Scleranthopsis within Acanthophyllum s.l. was confirmed by previous molecular studies, while the positions of Diaphanoptera and Kuhitangia remained uncertain. We have performed an updated molecular study of Acanthophyllum s.l. including an increased sampling of the genera and sections assigned to this group, using intron sequences of the chloroplast gene rps16 and nuclear ribosomal internal transcribed spacer (ITS) sequences. Cyathophylla, Heterochroa, and Saponaria were chosen as outgroups for performing phylogenetic analyses using maximum likelihood and Bayesian methods. The present results suggest that, in addition to the genera mentioned above, both Diaphanoptera and Kuhitangia should also be synonymized within Acanthophyllum. Sections Diaphanoptera, Kuhitangia and Pseudomacrostegia are introduced as new infrageneric taxa within Acanthophyllum. Our results also indicate that some annual species of Saponaria are closely related to Acanthophyllum.

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Acanthophyllum Meyer (1831: 210) in a broad sense comprises 80–90 perennial subshrubby species which are distributed mainly in the Irano-Turanian region (see e.g., Bittrich 1993, Ghaffari 2004, Pirani et al. 2014). According to the phylogenetic study by Pirani et al. (2014), Acanthophyllum s.lat. includes 11 sections. As part of our ongoing taxonomic investigations of Acanthophyllum, corrections to names of five Acanthophyllum sections are made here.
Chapter
Annual or perennial herbs, or subshrubs, rarely shrubs or small trees, usually monoecious, rarely gynodioecious or dioecious. Stems often swollen at the nodes, usually with anomalous secondary growth in older stems, also often occurring in the roots. Leaves opposite, decussate or apparently whorled, very rarely alternate, simple, entire, often connate at the base, sometimes succulent; stipules wanting, more rarely present, then mostly scarious. Flowers commonly protandrous, actinomorphic, very rarely weakly zygomorphic, usually bisexual, rarely unisexual, hypogynous or perigynous, sometimes obdiplostemonous. Inflorescences mostly dichasial, sometimes monochasial, rarely flowers solitary. Sepals (4−)5 or rarely more or fewer, imbricate, rarely valvate, completely free or connate for often most of their length, sometimes subtended by bracts (epicalyx). Petals (4−)5 or rarely more, free, aestivation contorted or rarely imbricate, entire, emarginate, bifid or lacerate, clawed or not, sometimes with coronal scales at the top of the claw, occasionally very small or absent.
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This work was undertaken at the suggestion of Dr. J. Lanjouw and Dr. F. P. Jonker. The citation heading this paragraph indicates that the group of Caryophyllaceae with which it deals, presents unusual taxonomic difficulties. At first, it was intended to restrict the revision to the genus Gypsophila. However, in the course of the work it was realized that the small genera Bolanthus, Ankyropetalum and Phryna could not be left out of account as they had been regarded by some authors as subdivisions of Gypsophila and by others as near relatives of this genus. For this reason a complete revision of these genera too was included. The only previous revision of Gypsophila is that published by Williams (1889). His study, largely based on data derived from the literature, includes 76 species i.e. about 3/5th of the number recognized here. His views on the generic limits were strongly influenced by those expressed by BENTHAM in BENTHAM and HOOKER’s Genera Plantarum 1 (1862). Later authors did not follow him in this respect, and generally preferred BOISSIER’s delimitation (1867), so e.g. PAX and HOFFMANN in the 2nd Edition of ENGLER und PRANTL, Planzenfam. (1934). PAX had already accepted this delimitation in the first edition (1889).
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The shape and the mode of dehiscence of the capsule had been regarded as good differential characters betweenAcanthophyllum and related genera.—Studies of these characters, including the shape of the ovary, in species ofAcanthophyllum, Diaphanoptera, Ochotonophila andScleranthopsis show, however, that they cannot be used as differential characters for the genusAcanthophyllum.
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Embryo, when differentiated, always with one cotyledon. The cotyledon usually with two main vascular bundles. Leaf venation striate or of derived types, mostly arcuate-striate or longitudinally striate (parallel), less often palmate-striate or pinnate-striate, almost always more or less closed at the apex (the veins emerging from the leaf base usually run together again at their apices). Leaves usually not clearly divided into petiole and lamina, less often more or less differentiated, but in these cases the “petiole” and the “lamina” are not homologous to those of magnoliopsids (are of secondary origin), often with sheathing base. Leaf traces usually numerous. Prophylls (including bracteoles) usually solitary and nearly always adaxial. Vascular bundles usually without cambium or rarely with vestigial cambium only. Vascular system of the stem usually consists of many separate scattered bundles or sometimes of two or more rings of vascular bundles, and the axis mostly attains its full diameter early, after which no increase in thickness takes place; only in some groups does thickening of the axis occur by means of division and enlargement of ground parenchyma cells (so-called diffuse secondary growth), as in palms, or by means of special kind of cambium that arises in the parenchyma outside the primary vascular system, as in some herbaceous and woody Lilianae. Sieve-element plastids of P-type with several to numerous cuneate (triangular) crystalloid bodies (lacking in all magnoliopsids studied except Saruma and Asarum in Aristolochiaceae). Phloem without parenchyma. Usually without clearly differentiated bark and pith. The primary root is usually ephemeral, dries out early in the growth of the plant, and is replaced by an adventitious root system that develops from the stem or (as in grasses) directly from the hypocotyl. Ontogenetically root cap and root epidermis are of different origin. Usually herbs, but often secondarily arborescent plants (primary woody plants are absent among the monocots). Flowers usually 3-merous, sometimes 4- or 2-merous, very rarely 5-merous. Nectaries predominantly septal. Pollen grains mostly 1-colpate (sulcate) or of derived types, often 1-porate.
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Fourteen polymorphic microsatellite loci in the endangered fern Adiantum reniforme var. sinense were developed and characterized using the Fast Isolation by AFLP of Sequences Containing repeats (FIASCO) protocol. Polymorphism of each locus was assessed in a bulked sample of 30 individuals from 8 natural populations. The number of alleles per locus varied from 2 to 11, with an average value of 6.2. The ranges of observed and expected heterozygosity were 0.000–0.895 and 0.226–0.868, respectively. These microsatellite markers provide useful tools for the ongoing conservation genetic studies of Adiantum reniforme var. sinense.
Article
A highly conserved sequence was found in rRNA gene internal transcribed spacer 1 (ITS1) among flowering plant species. The sequence, GGCRY-(4 to 7 n)-GYGYCAAGGAA (where Y = C or T; R = G or A) is located in the central region of ITS1, and is present in published sequences from a wide range of flowering plants. The rest of ITS1 is highly variable in sequence. Therefore, the conserved motif within ITS1 may have a key function in the processing of rRNA gene transcripts. Furthermore, identification of such a conserved motif will help facilitate alignment of sequences for phylogenetic analysis.
Alexey Grebenjuk (Komarov Botanical Institute of the Russian Academy of Sciences
  • B M Fumh
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  • Tari Msb
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  • Wu
HM, http://orcid.org/0000-0002-2406-2666, hmoazzeni@um.ac.ir; SZ, https://orcid.org/0000-0001-9159-1800, zarre@khayam.ut.ac.ir; RKR, https://orcid.org/0000-0002-6765-0353, rabeler@umich.edu; BO, https://orcid.org/0000-0002-6104-4264, bengt.oxelman@bioenv.gu.se; AVP, https://orcid.org/0000-0003-2494-0702, alexpavlenko1974@gmail.com; AK, https://orcid.org/ 0000-0001-8704-4644, andriy.kovalchuk@helsinki.fi. ■ ACKNOWLEDGMENTS We are grateful to Mohammad Mahmoodi (Research Institute of Forests and Rangelands, Tehran), Jalil Noroozi (University of Vienna, Vienna), Saeide Hoseini (Ferdowsi University of Mashhad, Mashhad), Farshid Memariani (Ferdowsi University of Mashhad, Mashhad), Alexey Grebenjuk (Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg), Aleksandr Ebel (Tomsk State University, Tomsk), Alim Gaziev (Tashkent), and Georgy Lazkov (Institute for Biology, National Academy of Sciences, Bishkek), for their kind help during different steps of this work. We also wish to thank curators at B, BM, FUMH, G, GB, H, JE, K, LD, LE, M, MSB, TARI, TASH, TK, TMRC, TUH and WU
Conspectus florae Asiae Mediae
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Bondarenko, O.N. 1971. Sem, Caryophyllaceae -Gvozdichnye [Family Caryophyllaceae] (pro parte). Pp. 225-310 in: Vvedensky, A.I. & Kovalevskaja, S.S. (eds.), Conspectus florae Asiae Mediae, vol. 2. Tashkent: Editio Academiae Scientarum UzSSR. [in Russian].
Saponaria Kermanensis Bornm. Nov. spec. sectionis Proteiniae e flora Persiae austro‐orientalis
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Verzeichniss der Arten der Gattung Acanthophyllum
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Notes on Caryophyllaceae, 3. On the genus Psammophiliella Ikonn
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Novyj rod iz semejstva Gvozdichnyh Pamiro‐Alaya – Kuhitangia (Kuhitangia Ovcz., gen. nov.) [A new genus of family Caryophyllaceae from Pamiro‐Alay – Kuhitangia (Kuhitangia Ovcz., gen. nov.)]
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Acanthophyllum transhyrcanum
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De quibusdam plantis novis Asiae mediae e familia Caryophyllacearum
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Flora of the U.S.S.R
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Flora Tadzhikskoi SSR
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