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Mary E. Endress, Raymond W. J. M. van der Ham, Siwert Nilsson, Laure Civeyrel, Mark W. Chase, Bengt Sennblad, Kurt Potgieter, Jeffrey Joseph, Martyn Powell, David Lorence, Ylva-Maria Zimmerman, Victor A. Albert
Within Rauvolfioideae (Apocynaceae), genera have long been assigned to tribes based mainly on only one or two superficial fruit and seed characters. Taxa with drupaceous fruits were included in Alyxieae. To elucidate relationships within Alyxieae, we analyzed phylogenetically a data set of sequences from four plastid DNA regions (RbcL, matK, TrnL intron, and TrnL-F intergenic spacer) and a morphological data set for 33 genera of Apocynaceae, including representatives of all genera previously included in Alyxieae and two non-Apocynaceae species. Results of parsimony analysis indicate that Alyxieae as previously delimited are polyphyletic, with most genera falling into two main clades. The Alyxia clade includes seven genera: Alyxia Banks ex R. Br., Lepinia Decne., Lepiniopsis Valeton, Pteralyxia K. Schum., and Condylocarpon Desf. together with Plectaneia Thouars. (earlier included in Plumerieae) and Chilocarpus Blume (earlier included in Chilocarpeae). The Vinca clade includes eight genera: Cabucala Pichon, Petchia Livera, Rauvolfia L., Catharanthus G. Don, Vinca L., Neisosperma Raf., Ochrosia Juss., and Kopsia Blume. Vallesia Ruiz & Pav. and Anechites Griseb. are not related to either clade and come out as sister to Aspidosperma Mart. & Zucc. (Aspidospermeae) and Thevetia L. (Plumerieae), respectively. The fruit and seed characters previously used to demarcate Alyxieae are homoplasious, as are other morphological characters such as style head structure and syncarpy versus apocarpy. Conversely, pollen morphology, which has not previously played much of a role in tribal delimitation, was shown to be the most useful morphological character for delimiting Alyxieae from other tribes of Rauvolfioideae.
The monotypic genera Absolmsia Kuntze, Madangia P. I. Forst., Liddle & I. M. Liddle, and Micholitzia N. E. Br. (Apocynaceae, Marsdenieae) are, according to molecular phylogenetic results, nested within the genus Hoya R. Br. Together with Dischidia R. Br., these genera are considered to comprise a well-supported clade within the tribe Marsdenieae. Flower morphology of Absolmsia,Madangia, and Micholitzia is examined by SEM and light microscopy and compared to that of the related H. caudata Hook. f., H. curtisii King & Gamble, H. hypolasia Schltr., H. lacunosa Blume, H. patella Schltr., H. retusa Dalzell, and H. venusta Schltr. We discuss the morphological evidence that supports the reduction of these genera into Hoya. Among characters supporting such a position is the presence of a pellucid margin on the pollinia of Absolmsia, Madangia, Micholitzia, and Hoya that is absent on the pollinia of Dischidia. Absolmsia and Madangia have a staminal corona with outer and inner lobes, similar to that of Hoya. While the outer corona processes of Absolmsia and Micholitzia are free and revolute, as in most species of Hoya, those of Madangia are laterally fused forming a continuous skirt resembling the corona of certain species of Hoya. The flowers of Micholitzia are superficially similar to those of Dischidia in its urceolate corolla, but the staminal corona with revolute outer processes decidedly links Micholitzia to Hoya. The new combination H. inflata (P. I. Forst., Liddle & I. M. Liddle) L. Wanntorp & P. I. Forst. is proposed. For Micholitzia obcordata N. E. Br., the name H. yuennanensis Hand.-Mazz. has nomenclatural priority and should therefore be used when including M. obcordata in Hoya.
Sequence data from the nuclear ITS region of 118 species (152 accessions) were used to test the monophyletic status and interrelationships of four related tribes in the Brassicaceae: Anchonieae, Chorisporeae, Euclidieae, and Hesperideae. Both maximum parsimony and maximum likelihood analyses support the recognition of the tribes Hesperideae (unigeneric, Hesperis L.) and Chorisporeae (3 genera, including Chorispora R. Br. ex DC., Diptychocarpus Trautv., and Parrya R. Br.), whereas the Anchonieae and the Euclidieae each separate into two distinct and distant clades (designated here as Anchonieae I and II and Euclidieae I and II). The data also support the exclusion of eight genera from the latter four tribes, with appropriate tribal assignment given in parentheses: Aubrieta Adans. (Arabideae), Blennodia R. Br. (Camelineae), Erysimum L. (Camelineae), Goldbachia DC. (unresolved), Hesperidanthus (B. L. Rob.) Rydb. (Schizopetaleae), Notothlaspi Hook. f. (unresolved), Pseudocamelina (Boiss.) N. Busch (Thlaspideae), and Zuvanda (Dvorák) Askerova (unresolved). The genus Malcolmia R. Br. (ca. 35 species) was paraphyletic and divided into three tribal clades, providing support for the separate genera Malcolmia, Strigosella Boiss., and Zuvanda. The genera Anchonium DC., Desideria Pamp., Eremobium Boiss., Erysimum, Iskandera N. Busch, Matthiola R. Br., Morettia DC., Neotorularia Hedge & J. Léonard, Rhammatophyllum O. E. Schulz, Sisymbriopsis Botsch. & Tzvelev, Sterigmostemum M. Bieb., and Tetracme Bunge each fell within a single tribal clade but were not monophyletic.
Ainsliaea DC. is an Asiatic genus of Mutisieae Cass. Forty-nine species and 19 varieties are recognized. Twenty-six species are endemic to China, four to Japan, four to Indo-China, two to the Himalayas, and one to Taiwan. The genus is characterized by its stems unbranched, with leaves basally rosulate or congested at the median part of the stem (only one species has stems branched, with alternate leaves), capitula few-flowered, florets deeply 5-cleft with unilateral lobes, and plumose pappus. Three new combinations are made: Ainsliaea bonatii Beauverd var. multibracteata (Mattf.) S. E. Freire, A. glabra Hemsl. var. sutchuenensis (Franch.) S. E. Freire, and A. henryi Diels var. subalpina (Hand.-Mazz.) S. E. Freire. One variety, A. latifolia (D. Don) Sch. Bip. var. taiwanensis S. E. Freire is described for the first time. The following taxa are newly synonymized: A. macroclinidioides Hayata var. secundiflora (Hayata) Kitam. and A. secundiflora Hayata with A. apiculata Sch. Bip. var. acerifolia Masam.; A. aptera DC. f. paniculata Kuntze with A. aptera DC.; A. pteropoda DC. var. platyphylla Franch. with A. bonatii Beauverd; A. mattfeldiana Hand.-Mazz. with A. bonatii var. multibracteata; A. cleistogama C. C. Chang with A. cavaleriei H. Lév.; A. asarifolia Hayata, A. cordifolia Franch. & Sav. var. integrifolia Maxim., A. fragrans Champ. ex Benth. var. integrifolia (Maxim.) Kitam., A. integrifolia (Maxim.) Makino, A. ningpoensis Matsuda, and A. rubrifolia Franch. with A. fragrans Champ. ex Benth.; A. lancifolia Franch. with A. glabra Hemsl.; A. tenuicaulis Mattf., A. glabra var. tenuicaulis (Mattf.) C. C. Chang, and A. plantaginifolia Mattf. with A. glabra Hemsl. var. sutchuenensis (Franch.) S. E. Freire; A. angustifolia Hook. f. & Thoms. ex C. B. Clarke with A. glumacea (Fr.) Sch. Bip.; A. gracilis Franch. var. robusta Diels with A. grossedentata Franch.; A. morrisonicola Hayata, A. reflexa Merr. var. morrisonicola (Hayata) Yamam., A. henryi Diels var. ovatifolia C. C. Chang, A. reflexa Merr. var. nimborum Hand.-Mazz., and
This is a study of the comparative foliar morphology carried out in seven neotropical species of the genus Marattia Sw.: M. alata Sw., M. cicutifolia Kaulf., M. excavata Underw., M. interposita Christ, M. laevis Sm., M. laxa Kunze, and M. weinmanniifolia Liebm., with the purpose of analyzing their diagnostic characters. A key to identify the species is obtained from the results.
Foliar characters were studied in 25 European, North African, North American, and South American terrestrial, amphibious, and aquatic taxa of the genus Isoetes L. (Pteridophyta, Isoetaceae). The foliar characters investigated have a variable diagnostic value. The presence and types of cuticular ornamentations, types of margins, epidermal patterns and cell ratios of the epidermis, types of IPP (intercellular pectic protuberances) in the cells of the leaf diaphragms, epidermal patterns and cell ratios of velum, epidermal patterns, and nature of walls in the adaxial epidermis of sporangia may be good diagnostic specific characters that should be included in taxonomic descriptions. Transverse sections of the microphylls, the types and distribution of collenchyma (fascicular, lacunar, or partially laminar collenchyma, with the latter two types described here for the first time), the cell types of leaf diaphragms, the presence of IPP in these cells, the partial or complete endodermis in the air chambers, and the presence of a persistent mucilage-secreting or an early lignified ligule are features that characterize species groups in Isoetes. The mucilage-secreting ligula and the types of endodermis can be considered as generic traits in Isoetes. The genus was largely considered as lacking good diagnostic characters or having rather environmentally influenced traits, two concepts that are discussed here after presentation of the new evidence.
We studied the floral biology of five North American members of Oenothera L. sect. Lavauxia (Spach) Endl. (Onagraceae L.) in field and common greenhouse settings. Oenothera sect. Lavauxia floral morphology ranges from small, cleistogamous flowers (O. flava subsp. flava (A. Nels.) Garrett in Garrett) to some of the longest-tubed flowers in North America (O. flava subsp. taraxacoides (Wooton & Standl.) W. L. Wagner). Our goal was to compare qualitative and quantitative aspects of floral advertisement and reward among taxa in section Lavauxia. All taxa are night-blooming and self-compatible, have yellow petals with ultraviolet reflectance, and produce floral scents dominated by nitrogenous compounds and monoterpenes. Methyl nicotinate is present in the fragrances of all taxa of section Lavauxia regardless of flower size or putative mating system. Because this floral volatile is largely absent from other Oenothera species, we hypothesize that it is a synapomorphy for section Lavauxia. The rare O. acutissima W. L. Wagner, which is endemic to the Uintah Mountains, is polymorphic for odors dominated by linalool- or ocimene-derived compounds. Field observations in its type locality in northeastern Utah, U.S.A., revealed frequent floral visitation by crepuscular hawkmoths during the first 1.5 hours after anthesis, a pattern common to O. flava subsp. taraxacoides and other large-flowered Oenothera throughout western North America. Quantitative aspects of floral advertisement (flower size, scent emission) and reward (nectar volume) are dramatically reduced in putatively autogamous taxa (O. flava subsp. flava, O. triloba Nutt.), whereas qualitative aspects (flower color, scent, and nectar chemistry) remain comparable. All taxa could be distinguished through ordination of characters related to flower size, herkogamy, and scent chemistry. Extreme nectar tube length variation across the range of O. flava renders this an excellent model system for measuring the costs and mechanisms of shifts between outcrossing and autogamy.
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