DOCSLIB.ORG

Cretaceous Asterid Evolution: Fruits of Eydeia Jerseyensis Sp

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Cretaceous Asterid Evolution: Fruits of Eydeia Jerseyensis Sp Annals of Botany 123: 451–460, 2019 doi: 10.1093/aob/mcy170, available online at www.academic.oup.com/aob Cretaceous asterid evolution: fruits of Eydeia jerseyensis sp. nov. (Cornales) from the upper Turonian of eastern North America Brian A. Atkinson1,2,*, Camila Martínez3 and William L. Crepet3 1Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66044, USA, 2Biodiversity Institute and Museum of Natural History, University of Kansas, Lawrence, KS 66044, USA and 3L. H. Bailey Hortorium, Plant Biology Downloaded from https://academic.oup.com/aob/article/123/3/451/5096126 by guest on 28 September 2021 Section, School of Integrative Plant Science, Cornell University, Ithaca, NY 14853, USA *For correspondence. E-mail [email protected] Received: 21 March 2018 Returned for revision: 5 May 2018 Editorial decision: 13 August 2018 Accepted: 10 September 2018 Published electronically 13 September 2018 • Background and aims The asterids (>80 000 extant species) appear in the fossil record with considerable diversity near the Turonian–Coniacian boundary (~90 Ma; Late Cretaceous) and are strongly represented in the earliest diverging lineage, Cornales. These early asterid representatives have so far been reported from western North America and eastern Asia. In this study, we characterize a new cornalean taxon based on charcoalified fruits from the upper Turonian of eastern North America, a separate landmass from western North America at the time, and identify early palaeobiogeographical patterns of Cornales during the Cretaceous. • Methods Fossils were studied and imaged using scanning electron microscopy and micro-computed tomogra- phy (micro-CT) scanning. To assess the systematic affinities of the fossils, phylogenetic analyses were conducted using maximum parsimony. • Key results The charcoalified fruits are represented by tri-locular woody endocarps with dorsal apically open- ing germination valves. Three septa intersect to form a robust central axis. Endocarp ground tissue consists of two zones: an outer endocarp composed of isodiametric sclereids and an inner endocarp containing circum-locular fibres. Central vasculature is absent; however, there are several small vascular bundles scattered within the septa. Phylogenetic analysis places the new taxon within the extinct genus Eydeia. • Discussion Thick-walled endocarps with apically opening germination valves, no central vascular bundle and one seed per locule are indicative of the order Cornales. Comparative analysis suggests that the fossils represent a new species, Eydeia jerseyensis sp. nov. This new taxon is the first evidence of Cornales in eastern North America during the Cretaceous and provides insights into the palaeobiogeography and initial diversification of the order. Key words: Asterids, Cornales, core eudicots, Cretaceous, fossils, fruits, New Jersey, palaeobiogeography. INTRODUCTION charcoalified flowers, an abundance of ericalean taxa have been characterized from the uppermost Turonian of eastern Exceptionally well-preserved fossils recently recovered from North America (Nixon and Crepet, 1993; Crepet et al., 2013; Upper Cretaceous deposits (~100–66 Ma) have provided mean- Martínez et al., 2016). For much of the Late Cretaceous, North ingful advancements in our understanding of the initial diversifi- America was divided by the Western Interior Seaway, sepa- cation of major core eudicot clades, the asterids and rosids (e.g. rating two distinct landmasses, Laramidia (western half) and Friis et al., 2011, 2016; Atkinson, 2016, 2018; Martínez et al., Appalachia (eastern half). Abundant palaeontological data sug- 2016; Atkinson et al., 2018). The earliest known core eudicots gest that northern Laramidia was connected with eastern Asia are rosid flowers and infructescences from the middle Cretaceous during the Late Cretaceous (discussed in Farke et al., 2014). (~100 Ma) of North America (Basinger and Dilcher, 1984; Friis However, despite their strong representation at the Turonian– et al., 2016; Manchester et al., 2018). Asterids appear relatively Coniacian boundary, Cornales have not yet been recovered late in the fossil record, near the Turonian–Coniacian boundary from Appalachia and Ericales have not yet been recovered from (~90–89 Ma) (Takahashi et al., 2002; Martínez-Millán, 2010; Laramidia or Asia (see Atkinson et al., 2018 for a discussion Friis et al., 2011; Manchester et al., 2015; Atkinson et al., 2018), on early asterids). It appears that each order initially diversified indicating that the initial phase of core eudicot evolution was in separate geographical regions, suggesting early endemism. dominated by rosid taxa; however, this pattern could be due to In this study, we characterize a new asterid taxon based on reporting and sampling biases. 106 charcoalified fruits from the upper Turonian Old Crossman Turonian–Coniacian asterids are represented by the two Clay Pit locality. The morphology and anatomy of these fruits successive earliest-diverging lineages, Cornales and Ericales, are indicative of the order Cornales. Phylogenetic analyses respectively. A diversity of cornalean fruits are known from and their unique combination of characters indicates that they the lowermost Coniacian of Asia and western North America represent a new species, Eydeia jerseyensis sp. nov. Atkinson, (Takahashi et al., 2002; Atkinson et al., 2018). Based on Martínez et Crepet. This report corresponds to the first record © The Author(s) 2018. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: [email protected]. 452 Atkinson et al. — Cretaceous Cornales evolution of Cornales in Appalachia during the Cretaceous, and expands Facility, Corvallis, OR, USA). Micro-computed tomography their known geographical distribution at the time of their earli- (micro-CT) scanning was also used to view internal structure of est appearance in the fossil record. The early diversity and dis- the holotype fruit (CUPC 1601) and was conducted at Cornell tribution of the cornalean fossil record supports the hypothesis University on the Zeiss Versa 520, at 60 kV/5 W, using the ×4 that the asterids were diversifying alongside the rosids prior to objective and an LE3 filter. The data were reconstructed from the Coniacian (Atkinson et al., 2018). 1601 fluoroscopy exposures of 1 s. The voxel size was 3.71 µm. The program used to visualize the CT scan results was OsiriX 64 bit DICOM Viewer. Images were processed using Photoshop MATERIALS AND METHODS CS 5.0 (Adobe, San Jose, CA, USA). Palaeobiogeographical maps (Fig. 5) were obtained from Scotese (2001) and Colorado One hundred and six charcoalified fruits were recovered from Plateau Geosystems (2014). the Old Crossman Clay Pit locality, south of the Raritan River Downloaded from https://academic.oup.com/aob/article/123/3/451/5096126 by guest on 28 September 2021 in Sayreville, Middlesex County, NJ, USA (40°28’ N, 74°19′ W; Fig. 1). Geologically, this locality belongs to the South Phylogenetic analyses Amboy Fire Clay Member of the Raritan Formation (Fig. 1). The Raritan Formation is composed of four members, of which the With the addition of the taxon described in this study, the mor- upper one corresponds to the South Amboy Fire Clay Member phological data matrix generated by Atkinson (2018) using 59 (Christopher, 1979; Sirkin, 1991). This member consists of mas- taxa (25 fossil and 34 extant) and 77 characters was utilized as sive unconsolidated to laminated, dark grey clay, and contains the basis of the phylogenetic analyses using maximum parsimony lignitized plant fragments, charcoalified flowers, stems, wood and is available through MorphoBank (O’Leary and Kaufman, fragments, fruits, and small pieces of amber that may have inclu- 2012; Project P3236). Seven characters were deactivated to sions of plant or insect material (Sugarman et al., 2005; Grimaldi improve phylogenetic resolution (Atkinson, 2018). The matrix and Nascimbene, 2010). The unconsolidated clay-silt beds gener- was analysed with and without a constraint topology. For the con- ally occur in channels, and are commonly adjacent to cross-bed- straint topology six nodes, grouping modern clades known from ded, fine to medium quartz sands with thin carbonaceous layers molecular phylogenies (Xiang et al., 2011), were constrained: (Sugarman et al., 2005). One of the more prevalent hypotheses (1) Cornaceae + Alangiaceae; (2) Nyssaceae–Mastixiaceae– to explain the three-dimensional preservation at the cellular level Davidiaceae; (4) Hydrangeaceae; (5) Loasaceae; and (6) the of these charcoalified fossils is that wildfires converted plant Ericales outgroup. None of the fossil taxa was constrained for the remains to carbonized replicas that were buried in the forest leaf analysis. The phylogeny was estimated based on maximum par- litter (Grimaldi and Nascimbene, 2010; Friis et al., 2011). simony criteria using TNT (Goloboff, 1998) through Winclada The paucity of marine invertebrates in rocks from the version 1.99 (Nixon, 2015). In the TNT analyses, ten sets of 500 Cretaceous Atlantic Coastal Plain has prevented precise cor- iterations and 10 % perturbation of characters were used for the relations of the lithological units with zonations based on ratchet analysis (Nixon, 1999), and default values were used for marine organisms; therefore, age determinations have been drift, sectorial search and tree fusion. A strict consensus and a based primarily on palynology (Christopher,
Load more

Recommended publications
  • Department of Planning and Zoning
    Department of Planning and Zoning Subject: Howard County Landscape Manual Updates: Recommended Street Tree List (Appendix B) and Recommended Plant List (Appendix C) - Effective July 1, 2010 To: DLD Review Staff Homebuilders Committee From: Kent Sheubrooks, Acting Chief Division of Land Development Date: July 1, 2010 Purpose: The purpose of this policy memorandum is to update the Recommended Plant Lists presently contained in the Landscape Manual. The plant lists were created for the first edition of the Manual in 1993 before information was available about invasive qualities of certain recommended plants contained in those lists (Norway Maple, Bradford Pear, etc.). Additionally, diseases and pests have made some other plants undesirable (Ash, Austrian Pine, etc.). The Howard County General Plan 2000 and subsequent environmental and community planning publications such as the Route 1 and Route 40 Manuals and the Green Neighborhood Design Guidelines have promoted the desirability of using native plants in landscape plantings. Therefore, this policy seeks to update the Recommended Plant Lists by identifying invasive plant species and disease or pest ridden plants for their removal and prohibition from further planting in Howard County and to add other available native plants which have desirable characteristics for street tree or general landscape use for inclusion on the Recommended Plant Lists. Please note that a comprehensive review of the street tree and landscape tree lists were conducted for the purpose of this update, however, only
    [Show full text]
  • Quali-Quantitative Analysis of Flavonoids of Cornus Mas L
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Archivio della ricerca della Scuola Superiore Sant'Anna Food Chemistry 119 (2010) 1257–1261 Contents lists available at ScienceDirect Food Chemistry journal homepage: www.elsevier.com/locate/foodchem Analytical Methods Quali-quantitative analysis of flavonoids of Cornus mas L. (Cornaceae) fruits Agata Maria Pawlowska a, Fabiano Camangi b, Alessandra Braca a,* a Dipartimento di Scienze Farmaceutiche, Università di Pisa, Via Bonanno 33, 56126 Pisa, Italy b Scuola Superiore S. Anna di Studi, Universitari e di Perfezionamento di Pisa, Piazza Martiri della, Libertà 33, 56127 Pisa, Italy article info abstract Article history: The methanol extract obtained from the ripe fruits of Cornus mas L. (Cornaceae) have been phytochem- Received 2 April 2009 ically studied. On the basis of HPLC–PDA–MS/MSn analysis eight compounds have been identified as Received in revised form 10 June 2009 quercetin, kaempferol, and aromadendrin glycosilated derivatives. Three major compounds have been Accepted 31 July 2009 also isolated by Sephadex LH-20 column chromatography followed by HPLC and characterised by NMR spectroscopy. Moreover, LC–PDA–MS analysis of the red pigment revealed the presence of three antho- cyanins. The quantitative analysis of all compounds was reported. Keywords: Ó 2009 Elsevier Ltd. All rights reserved. Cornus mas Flavonoids Anthocyanins HPLC ESI-MS/MS 1. Introduction their subclass anthocyanins, contribute the yellow, orange, red, and blue colour to flowers, fruits, and vegetables (Cooper-Driver, Cornus mas L. (Cornaceae family) known as the European and 2001), and could become important in the replacing of the syn- Asiatic cornelian cherry is a species of dogwood native to Southern thetic pigments by the natural ones.
    [Show full text]
  • Papers in Press
    Papers in Press “Papers in Press” includes peer-reviewed, accepted manuscripts of research articles, reviews, and short notes to be published in Paleontological Research. They have not yet been copy edited and/or formatted in the publication style of Paleontological Research. As soon as they are printed, they will be removed from this website. Please note they can be cited using the year of online publication and the DOI, as follows: Humblet, M. and Iryu, Y. 2014: Pleistocene coral assemblages on Irabu-jima, South Ryukyu Islands, Japan. Paleontological Research, doi: 10.2517/2014PR020. doi:10.2517/2018PR013 Features and paleoecological significance of the shark fauna from the Upper Cretaceous Hinoshima Formation, Himenoura Group, Southwest Japan Accepted Naoshi Kitamura 4-8-7 Motoyama, Chuo-ku Kumamoto, Kumamoto 860-0821, Japan (e-mail: [email protected]) Abstract. The shark fauna of the Upper Cretaceous Hinoshima Formation (Santonian: 86.3–83.6 Ma) of the manuscriptHimenoura Group (Kamiamakusa, Kumamoto Prefecture, Kyushu, Japan) was investigated based on fossil shark teeth found at five localities: Himedo Park, Kugushima, Wadanohana, Higashiura, and Kotorigoe. A detailed geological survey and taxonomic analysis was undertaken, and the habitat, depositional environment, and associated mollusks of each locality were considered in the context of previous studies. Twenty-one species, 15 genera, 11 families, and 6 orders of fossil sharks are recognized from the localities. This assemblage is more diverse than has previously been reported for Japan, and Lamniformes and Hexanchiformes were abundant. Three categories of shark fauna are recognized: a coastal region (Himedo Park; probably a breeding site), the coast to the open sea (Kugushima and Wadanohana), and bottom-dwelling or near-seafloor fauna (Kugushima, Wadanohana, Higashiura, and Kotorigoe).
    [Show full text]
  • Cornus Kousa Kousa Dogwood1 Edward F
    Fact Sheet ST-191 November 1993 Cornus kousa Kousa Dogwood1 Edward F. Gilman and Dennis G. Watson2 INTRODUCTION Kousa Dogwood grows 15 to 20 feet tall and has beautiful exfoliating bark, long lasting flowers, good fall color, and attractive fruit (Fig. 1). Branches grow upright when the tree is young, but appear in horizontal layers on mature trees. The crown eventually grows wider than it is tall on many specimens. It would be difficult to use too many Kousa Dogwoods. The white, pointed bracts are produced a month later than Flowering Dogwood and are effective for about a month, sometimes longer. The red fruits are edible and they look like a big round raspberry. Birds devour the fruit quickly. Fall color varies from dull red to maroon. GENERAL INFORMATION Scientific name: Cornus kousa Pronunciation: KOR-nus KOO-suh Common name(s): Kousa Dogwood, Chinese Dogwood, Japanese Dogwood Family: Cornaceae USDA hardiness zones: 5 through 8 (Fig. 2) Figure 1. Young Kousa Dogwood. Origin: not native to North America Uses: container or above-ground planter; near a deck DESCRIPTION or patio; screen; specimen Availability: generally available in many areas within Height: 15 to 20 feet its hardiness range Spread: 15 to 20 feet Crown uniformity: symmetrical canopy with a regular (or smooth) outline, and individuals have more or less identical crown forms Crown shape: round Crown density: dense Growth rate: slow 1. This document is adapted from Fact Sheet ST-191, a series of the Environmental Horticulture Department, Florida Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida.
    [Show full text]
  • Tagawa Gardens June Snow Giant Dogwood
    June Snow Giant Dogwood Cornus controversa 'June Snow-JFS' Height: 30 feet Spread: 40 feet Sunlight: Hardiness Zone: 4 Description: June Snow Giant Dogwood flowers Covered with lovely white flower clusters in spring; a Photo courtesy of NetPS Plant Finder large rounded form, with gracefully layered branches make this tree an excellent specimen; attractive fruit in late summer turns deep blue-black; orange to red fall color Ornamental Features June Snow Giant Dogwood features showy clusters of white flowers held atop the branches in late spring. It has dark green foliage throughout the season. The pointy leaves turn an outstanding dark red in the fall. It produces navy blue berries from early to late fall. The warty gray bark and green branches add an interesting dimension to the landscape. June Snow Giant Dogwood in bloom Landscape Attributes Photo courtesy of NetPS Plant Finder June Snow Giant Dogwood is a deciduous tree with a stunning habit of growth which features almost oriental horizontally-tiered branches. Its average texture blends into the landscape, but can be balanced by one or two finer or coarser trees or shrubs for an effective composition. This is a relatively low maintenance tree, and should only be pruned after flowering to avoid removing any of the current season's flowers. It is a good choice for attracting birds to your yard. It has no significant negative characteristics. June Snow Giant Dogwood is recommended for the following landscape applications; - Accent - Shade Planting & Growing June Snow Giant Dogwood will grow to be about 30 feet tall at maturity, with a spread of 40 feet.
    [Show full text]
  • Steciana Doi:10.12657/Steciana.020.004 ISSN 1689-653X
    2016, Vol. 20(1): 21–32 Steciana doi:10.12657/steciana.020.004 www.up.poznan.pl/steciana ISSN 1689-653X ANATOMICAL STUDY OF CORNUS MAS L. AND CORNUS OFFICINALIS SEIB. ET ZUCC. (CORNACEAE) ENDOCARPS DURING THEIR DEVELOPMENT MARIA MOROZOWSKA, ILONA WYSAKOWSKA M. Morozowska, I. Wysakowska, Department of Botany, Poznań University of Life Sciences, Wojska Polskiego 71 C, 60-625 Poznań, Poland, e-mail: [email protected], [email protected] (Received: June 17, 2015. Accepted: October 20, 2015) ABSTRACT. Results of anatomical studies on the developing endocarps of Cornus mas and C. officinalis are pre- sented. Formation of an endocarp and anatomical changes in its structure from the flowering stage to fully developed fruits were observed with the use of LM and SEM. In the process of anatomical development of endocarps the formation of two layers, i.e. the inner and the outer endocarp, was observed. Changes in their anatomical structure consisted in a gradual thickening of cell walls and their lignification. The ligni- fication of endocarp cell walls begins in the inner endocarp, it proceeds in the outside parts of the outer endocarp, with an exception of several layers of cells forming the transition zone (circular strand) present on its margin, and finally, almost at the same time, in the rest of the outer endocarp. Cell walls of the cells distinguishing the germinating valves undergo thickening delayed by several days in comparison to other endocarp cells. Thickening of their cell walls starts in cells situated close to the inner endocarp and proceeds to its outer parts, and their lignification is not very intensive.
    [Show full text]
  • Cenomanian Turonian Coniacian Santonian Campanian
    walteri aff. aff. spp. spp. imperfectus spp. (prisms) Chronostratigraphy Offshore Norway sp. 1 Geologic Time Scale 2012 Zonation (Gradstein et al., 1999, and this study) Allomorphina halli / pyriformis Sigmoilina antiqua Textularia Gavelinella intermeda gracillima Valvulineria Bulbobaculites problematicus Caudammina ovuloides Nuttallinella florealis Stensioeina granulata polonica Inoceramus Rzehakina minima Rzehakina epigona Fenestrella bellii Gaudryina filiformis Trochamminoides Haplophragmoides Gavelinella usakensis Caudammina ovula Coarse agglutinated spp. LCO dubia Tritaxia Plectorecurvoides alternans Reussella szajnochae Recurvoides Hippocrepina depressa Psammosphaera sphaerical radiolarians Ma Age/Stage Lingulogavelinella jarzevae elegans Lt NCF19 Maastrichtian volutus LCO 70 NCF18 E szajnochae dubia Lt 75 LCO of NCF17 Campanian Deep Water M Agglutinated 80 Foraminifera E bellii NCF16 Lt Inoceramus LCO NCF15 85 Santonian M E polonica NCF14 Lt Coniacian M E Marginotruncana NCF13 90 Lt Turonian M E Dicarinella NCF12 95 Lt brittonensis M NCF11 Cenomanian delrioensis LCO NCF10 E antiqua NCF9 100 Figure 2.8c. Stratigraphic ranges of Upper Cretaceous benthic foraminifera, and miscellaneous index taxa, oshore mid-Norway, with the foraminiferal zonation established in this study. s.l. Chronostratigraphy Offshore Norway Geologic Time Scale 2012 Zonation (Gradstein et al., 1999, and this study) Abathomphalus mayaroensis Pseudotextularia elegans Hedbergella planispira Hedbergella hoelzi Praeglobotruncana delrioensis Praeglobotruncana stephani
    [Show full text]
  • From the Late Eocene of Hordle, Southern England
    Acta Palaeobotanica 59(1): 51–67, 2019 e-ISSN 2082-0259 DOI: 10.2478/acpa-2019-0006 ISSN 0001-6594 Fruit morphology, anatomy and relationships of the type species of Mastixicarpum and Eomastixia (Cornales) from the late Eocene of Hordle, southern England STEVEN R. MANCHESTER1* and MARGARET E. COLLINSON2 1 Florida Museum of Natural History, Dickinson Hall, P.O. Box 117800, Gainesville, Florida, U.S.A.; e-mail: [email protected] 2 Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey TW20 0EX, United Kingdom Received 26 October 2018; accepted for publication 29 April 2019 ABSTRACT. The Mastixiaceae (Cornales) were more widespread and diverse in the Cenozoic than they are today. The fossil record includes fruits of both extant genera, Mastixia and Diplopanax, as well as several extinct genera. Two of the fossil genera, Eomastixia and Mastixicarpum, are prominent in the palaeobotanical literature, but concepts of their delimitation have varied with different authors. These genera, both based on species described 93 years ago by Marjorie Chandler from the late Eocene (Priabonian) Totland Bay Member of the Headon Hill Formation at Hordle, England, are nomenclaturally fundamental, because they were the first of a series of fos- sil mastixioid genera published from the European Cenozoic. In order to better understand the type species of Eomastixia and Mastixicarpum, we studied type specimens and topotypic material using x-ray tomography and scanning electron microscopy to supplement traditional methods of analysis, to improve our understanding of the morphology and anatomy of these fossils. Following comparisons with other fossil and modern taxa, we retain Mas- tixicarpum crassum Chandler rather than transferring it to the similar extant genus Diplopanax, and we retain Eomastixia bilocularis Chandler [=Eomastixia rugosa (Zenker) Chandler] and corroborate earlier conclusions that this species represents an extinct genus that is more closely related to Mastixia than to Diplopanax.
    [Show full text]
  • Phytogeographic Review of Vietnam and Adjacent Areas of Eastern Indochina L
    KOMAROVIA (2003) 3: 1–83 Saint Petersburg Phytogeographic review of Vietnam and adjacent areas of Eastern Indochina L. V. Averyanov, Phan Ke Loc, Nguyen Tien Hiep, D. K. Harder Leonid V. Averyanov, Herbarium, Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov str. 2, Saint Petersburg 197376, Russia E-mail: [email protected], [email protected] Phan Ke Loc, Department of Botany, Viet Nam National University, Hanoi, Viet Nam. E-mail: [email protected] Nguyen Tien Hiep, Institute of Ecology and Biological Resources of the National Centre for Natural Sciences and Technology of Viet Nam, Nghia Do, Cau Giay, Hanoi, Viet Nam. E-mail: [email protected] Dan K. Harder, Arboretum, University of California Santa Cruz, 1156 High Street, Santa Cruz, California 95064, U.S.A. E-mail: [email protected] The main phytogeographic regions within the eastern part of the Indochinese Peninsula are delimited on the basis of analysis of recent literature on geology, geomorphology and climatology of the region, as well as numerous recent literature information on phytogeography, flora and vegetation. The following six phytogeographic regions (at the rank of floristic province) are distinguished and outlined within eastern Indochina: Sikang-Yunnan Province, South Chinese Province, North Indochinese Province, Central Annamese Province, South Annamese Province and South Indochinese Province. Short descriptions of these floristic units are given along with analysis of their floristic relationships. Special floristic analysis and consideration are given to the Orchidaceae as the largest well-studied representative of the Indochinese flora. 1. Background The Socialist Republic of Vietnam, comprising the largest area in the eastern part of the Indochinese Peninsula, is situated along the southeastern margin of the Peninsula.
    [Show full text]
  • Taxonomy of Alangium Section Conostigma (Alangiaceae)
    Blumea 62, 2017: 29–46 ISSN (Online) 2212-1676 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE https://doi.org/10.3767/000651917X695164 Taxonomy of Alangium section Conostigma (Alangiaceae) W.J.J.O. de Wilde1, B.E.E. Duyfjes1 Key words Abstract Alangium Lam. sect. Conostigma Bloemb. is largely confined to the Malesian area and contains 19 species. By using the characters as found in the original species-descriptions, and with emphasizing the nature of Alangiaceae the indument of the leaf bud, the confusingly variable Alangium javanicum s.l.-complex could be dismantled for a Alangium sect. Conostigma great part. Four new species: A. ledermannii, A. pallens, A. plumbeum, and A. subcordatum, and two new varieties: Malesia A. meyeri var. macilentum and A. ebenaceum var. insignis are recognised. Two taxa are raised to specific status: new species A. nobile subsp. denudatum to A. denudatum and A. javanicum var. minahassicum to A. minahassicum. A key to taxonomy the species is presented, and the concerned taxa are enumerated, referenced, described and some are figured. Published on 14 February 2017 INTRODUCTION In the present paper we propose to re-instate the majority of names of species formerly described in this complex of A. ja- The section Conostigma is one of the four sections recognised vanicum s.l., but later on sunk into synonymy. Also, some new in Alangium by Bloembergen (1935, 1939). His division into taxa are proposed. Unfortunately, occasional specimens remain sections was later corroborated with anatomical and molecular difficult to determine, and some of the species accepted seem characters (Eyde 1968, Feng et al.
    [Show full text]
  • GEOLOGIC TIME SCALE V
    GSA GEOLOGIC TIME SCALE v. 4.0 CENOZOIC MESOZOIC PALEOZOIC PRECAMBRIAN MAGNETIC MAGNETIC BDY. AGE POLARITY PICKS AGE POLARITY PICKS AGE PICKS AGE . N PERIOD EPOCH AGE PERIOD EPOCH AGE PERIOD EPOCH AGE EON ERA PERIOD AGES (Ma) (Ma) (Ma) (Ma) (Ma) (Ma) (Ma) HIST HIST. ANOM. (Ma) ANOM. CHRON. CHRO HOLOCENE 1 C1 QUATER- 0.01 30 C30 66.0 541 CALABRIAN NARY PLEISTOCENE* 1.8 31 C31 MAASTRICHTIAN 252 2 C2 GELASIAN 70 CHANGHSINGIAN EDIACARAN 2.6 Lopin- 254 32 C32 72.1 635 2A C2A PIACENZIAN WUCHIAPINGIAN PLIOCENE 3.6 gian 33 260 260 3 ZANCLEAN CAPITANIAN NEOPRO- 5 C3 CAMPANIAN Guada- 265 750 CRYOGENIAN 5.3 80 C33 WORDIAN TEROZOIC 3A MESSINIAN LATE lupian 269 C3A 83.6 ROADIAN 272 850 7.2 SANTONIAN 4 KUNGURIAN C4 86.3 279 TONIAN CONIACIAN 280 4A Cisura- C4A TORTONIAN 90 89.8 1000 1000 PERMIAN ARTINSKIAN 10 5 TURONIAN lian C5 93.9 290 SAKMARIAN STENIAN 11.6 CENOMANIAN 296 SERRAVALLIAN 34 C34 ASSELIAN 299 5A 100 100 300 GZHELIAN 1200 C5A 13.8 LATE 304 KASIMOVIAN 307 1250 MESOPRO- 15 LANGHIAN ECTASIAN 5B C5B ALBIAN MIDDLE MOSCOVIAN 16.0 TEROZOIC 5C C5C 110 VANIAN 315 PENNSYL- 1400 EARLY 5D C5D MIOCENE 113 320 BASHKIRIAN 323 5E C5E NEOGENE BURDIGALIAN SERPUKHOVIAN 1500 CALYMMIAN 6 C6 APTIAN LATE 20 120 331 6A C6A 20.4 EARLY 1600 M0r 126 6B C6B AQUITANIAN M1 340 MIDDLE VISEAN MISSIS- M3 BARREMIAN SIPPIAN STATHERIAN C6C 23.0 6C 130 M5 CRETACEOUS 131 347 1750 HAUTERIVIAN 7 C7 CARBONIFEROUS EARLY TOURNAISIAN 1800 M10 134 25 7A C7A 359 8 C8 CHATTIAN VALANGINIAN M12 360 140 M14 139 FAMENNIAN OROSIRIAN 9 C9 M16 28.1 M18 BERRIASIAN 2000 PROTEROZOIC 10 C10 LATE
    [Show full text]
  • Cornaceae Dogwood Family
    Cornaceae dogwood family North-temperate shrubs or trees, the dogwoods have few herbaceous perennials amongst them. Page | 487 Inflorescence is a cyme, often subtended by showy bracts. Four or five-merous, stamens oppose the petals, and are of equal number, or totalling 15 arranged in whorls. Calyx may be present or absent, and may be reduced to a rim around the inferior ovary. Fruit is a drupe, the stone grooved longitudinally. Leaves are typically opposite and seldom alternate. Cornus dogwoods About 50 species are included here; three shrubs and two herbs reach Nova Scotia. Flowers are four- merous, their sepals minutes and petals small. Leaves have distinctive venation. Key to species A. Inflorescence an open cyme, bracts minute or absent; fruit maturing blue to B white; shrubs. B. Leaves alternate, clustered distally. Cornus alternifolia bb. Leaves opposite. C C. Twigs red; fruit white, stone dark brown with yellow C. sericea stripes. cc. Twigs not red; fruit blue to white, stone pale. C. rugosa aa. Inflorescence a dense head, subtended by 4 showy bracts; fruit maturing D bright red; herbaceous. D. Lateral veins arising from the midrib along the leaf. C. canadensis dd. Lateral veins arising only from the base of the leaf. C. suecica Cornus alternifolia L.f. Alternate-leaved Dogwood; cornouiller à feuilles alternes A shrub with alternate leaves, their margins are smooth. Leaves are clustered at the apices of the branches. Veins strongly mark the leaves, curving to the acute apices. Stems are yellow. Inflorescence is a round cyme of many creamy flowers, producing blue drupes. Flowers mid-June to mid-July.
    [Show full text]