From the Late Eocene of Hordle, Southern England

Total Page:16

File Type:pdf, Size:1020Kb

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. KEYWORDS: fruits, Mastixia, Diplopanax, fossil, Late Eocene, Tertiary INTRODUCTION Both extant genera of the cornalean family has been recognized (reviewed in Kirchheimer Mastixiaceae have distinctive endocarps that 1957, Mai 1993). are readily recognizable in the fossil record. Generic and specific nomenclature for fossil Although geographically limited to Asia mastixioid fruits is complicated because many (Diplopanax and Mastixia) and Malesia (Mas- of the fossil genera and species were named tixia) today, both genera are well documented before the fruit morphology and relationships by fossil fruits in the Cenozoic of Europe (Mai of the extant representatives were well under- 1970, Martinetto 2011, Hably & Erdei 2013) stood. The living genus Diplopanax Hand.- and North America (Tiffney & Haggard 1996, Mazz was named (Handel-Mazzetti 1933) sub- Stockey et al. 1998). In addition, a complex sequent to the designation of the extinct genera of extinct mastixioid genera with endocarps Mastixicarpum and Eomastixia by Chandler similar to those of Mastixia and Diplopanax (1926), but initially it was thought to repre- sent Araliaceae, and its affinities to Mastixia * Corresponding author were only recognized 50 years later (Zeng 1983, 52 S.R. Manchester & M.E. Collinson / Acta Palaeobotanica 59(1): 51–67, 2019 Eyde & Xiang 1990). Meanwhile, fossil fruits MATERIALS AND METHODS with similar morphology had been assigned to other genera such as Tectocarya Kirchheimer Specimens. Fruits of extant Mastixia and Diplo- (1936), Mastixiopsis Kirchheimer (1936), Mas- panax were borrowed from herbaria of the US National tixiodiocarpum Scott (Scott 1954, Manchester Museum, Washington, DC (US); Royal Botanic Gardens, 1994) and others. The recognition and distinc- Kew, Richmond, Surrey, UK (K); the Komarov Botani- tion of various fossil mastixioid genera were cal Institute, Saint Petersburg (LE), and the Missouri Botanical Garden, St. Louis (MO). Fossil specimens are informed by degrees of departure from what housed in the palaeobotany collections, Earth Depart- was then considered to be the only modern ment, Natural History Museum London (NHMUK; genus – Mastixia. Names proliferated, with up specimen numbers prefixed “V.”). Geological occurrence to 16 genera and 40 species in the literature on and locality details may be found in the original pub- lished sources (Chandler 1926, 1962, Reid & Chandler European fossil mastixioid fruits. Some were 1933), reviewed and updated by Collinson (1996) and subsequently synonomized, however, and Mai Cleal et al. (2001). The Hordle samples (Hordle bed 10) (1993) reduced the number of European fos- are from the Totland Bay Member of the Headon Hill sil mastixioid genera to seven, including five Formation of late Eocene (Priabonian) age. extinct genera plus fossil species of the two Light and scanning electron microscopy. Extant and fossil fruits were studied using natural frac- extant genera. Difficulties remain, however, ture surfaces and transverse sections cut with a paper- because some of the earliest-described taxa – thin diamond blade mounted on a Microslice II annular important because of their nomenclatural pri- saw. Naturally fractured surfaces were also studied by ority – are less well known anatomically than scanning electron microscopy (SEM) using a Hitachi some of the later-described examples. S-4000 instrument at the University of Florida. Micro-computed tomography (micro-CT). For Fossil mastixioid genera were first recog- anatomical investigation of extant material, selected nized by Chandler (1926) from the late Eocene fruits were examined by micro-CT at the Imaging and of Hordle, England. Based upon lignified fruits, Analysis Centre, NHMUK, using a Nikon Metrology she established two new genera that were sub- HMX ST 225. These included a fruit from the type sequently found to be widespread in the Euro- material of Mastixia pentandra (K704731) and another from type material of M. euonymoides (K704785), plus pean Tertiary: Eomastixia and Mastixicarpum. two other fruits of Mastixia euonymoides (G. Watt The circumscription of these genera was subse- 5899, K). To protect specimens and avoid using adhe- quently revised as new species from other sites sive, the fruits were first wrapped in cling film and were added (Kirchheimer 1957, Holý 1975, Mai then inserted carefully into Oasis TM (foam used in 1993), and by Chandler (1962) when she exam- flower arranging) for stability during the scan. Scan- ning was undertaken using a tungsten reflection target ined larger collections of topotypic material at accelerating voltage of 90 kV and current of 290 µA, from Hordle as well as material from other sites. with an exposure time of 708 ms (giving a scan time Chandler’s work, both in 1926 and 1962, was of 38 minutes). No filters were used and 3142 projec- careful and detailed, but limited by the tools of tions were taken over a 360° rotation. Voxel sizes of the time for imaging fine-structure details. Her the resulting datasets were 12 or 13 µm depending on specimen size. Three-dimensional volumes were descriptions were based on optical observations reconstructed using CT Pro (Nikon Metrology, Tring, of fruits in surface view and variously fractured UK) and TIFF stacks were exported using VG Studio to reveal internal anatomy of the fruit and seed Max (Volume Graphics GmbH, Heidelberg, Germany). features, and informed by careful comparative Datasets were visualized, and images and videos were work on available extant taxa. captured, using Avizo 8.1 (FEI Visualization Science Group, Bordeaux, France). Images were adjusted uni- For the present article we review the mor- formly overall for contrast and brightness using Adobe phology and anatomy of the extant Mastixia Photoshop® CS2 or CS6. Micro-CT scan datasets are and Diplopanax and provide detailed compara- archived at NHMUK. tive studies of the morphology and anatomy for Synchrotron radiation x-ray tomographic the type species of Eomastixa and Mastixicar- microscopy (SRXTM). For improved resolution of ana- tomical details, SRXTM was applied to selected fossil pum, based on fruits from the original locality specimens and fruits of extant Mastixia using the TOM- at Hordle, England. New anatomical details CAT beamline of the Swiss Light Source, Paul Scher- revealed by x-ray tomography and scanning rer Institut, Villigen, Switzerland, in March 2015 (fos- electron microscopy improve our understanding sils) and February 2016 (extant). For extant Mastixia, of the relationships among these modern and one fruit from type material of each of M. pentandra (K704731) and M. euonymoides (K704785) was scanned. fossil taxa, confirming that Eomastixia is close For fossil Mastixicarpum crassum specimens, the holo- to Mastixia and that Mastixicarpum is similar type V20074a [figured by Chandler 1962: pl. 19, fig. 19] to but distinct from Diplopanax. (detached valve in a separate scan from main fruit), S.R. Manchester & M.E. Collinson / Acta Palaeobotanica 59(1): 51–67, 2019 53 V68593i and V68593ii (two halves of the same speci- side of the endocarp, which becomes the ger- men, re-registered from V20076 duplicate material) mination valve. This is seen on the surface Eomas- from Hordle were scanned. Both syntypes of of the endocarp as a narrow longitudinal slit. tixia bilocularis (V20079a, V20079b, labeled E. rugosa in the collection) were scanned, as was V40768 from the This infold intrudes the locule along the plane Bournemouth Freshwater Beds, which was figured by of bisymmetry,
Recommended publications
  • Research Article
    Research Article: Maximum Likelihood Analyses of 3,490 rbcL Sequences: Scalability of Comprehensive Inference versus Group-Specific Taxon Sampling Alexandros Stamatakis1,*, Markus Göker2,3, Guido Grimm4 1 The Exelixis Lab, Dept. of Computer Science, Technische Universität München, Germany 2 Organismic Botany, Eberhard-Karls-University, Tübingen, Germany 3 German Collection of Microorganisms and Cell Cultures, Braunschweig, Germany 4 Department of Palaeobotany, Swedish Museum of Natural History, Stockholm, Sweden * Corresponding author: Technische Universität München, Fakultät für Informatik, I 12, Boltzmannstr. 3, 85748 Garching b. München, Tel: +49 89 28919434, Fax: +49 89 28919414, Email: [email protected] Keywords: Phylogenetic Inference, Maximum Likelihood, RAxML, large single–gene datasets, eudicots Running head: Scalability of Maximum Likelihood Analyses Abbreviations: GRTS: Group-based Randomized Taxon-Subsampling; TU: Taxonomic Unit. Abstract The constant accumulation of sequence data poses new computational and methodological challenges for phylogenetic inference, since multiple sequence alignments grow both in the horizontal (number of base pairs, phylogenomic alignments) as well as vertical (number of taxa) dimension. Put aside the ongoing controversial discussion about appropriate models, partitioning schemes, and assembly methods for phylogenomic alignments, coupled with the high computational cost to infer these, for many organismic groups, a sufficient number of taxa is often exclusively available from one or just a few genes (e.g., rbcL, matK, rDNA). In this paper we address scalability of Maximum-Likelihood-based phylogeny reconstruction with respect to the number of taxa by example of several large nested single-gene rbcL alignments comprising 400 up to 3,491 taxa. In order to thoroughly test the effect of taxon sampling, we deploy an appropriately adapted taxon jackknifing approach.
    [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]
  • Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa Grayensis Sp
    East Tennessee State University Digital Commons @ East Tennessee State University Electronic Theses and Dissertations Student Works 8-2013 Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee Nathan R. Noll East Tennessee State University Follow this and additional works at: https://dc.etsu.edu/etd Part of the Biodiversity Commons, Climate Commons, Paleontology Commons, and the Plant Biology Commons Recommended Citation Noll, Nathan R., "Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee" (2013). Electronic Theses and Dissertations. Paper 1204. https://dc.etsu.edu/etd/1204 This Thesis - Open Access is brought to you for free and open access by the Student Works at Digital Commons @ East Tennessee State University. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Digital Commons @ East Tennessee State University. For more information, please contact [email protected]. Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee ___________________________ A thesis presented to the faculty of the Department of Biological Sciences East Tennessee State University In partial fulfillment of the requirements for the degree Master of Science in Biology ___________________________ by Nathan R. Noll August 2013 ___________________________ Dr. Yu-Sheng (Christopher) Liu, Chair Dr. Tim McDowell Dr. Foster Levy Keywords: Nyssa, Endocarp, Gray Fossil Site, Miocene, Pliocene, Karst ABSTRACT Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp.
    [Show full text]
  • Report of Rapid Biodiversity Assessments at Cenwanglaoshan Nature Reserve, Northwest Guangxi, China, 1999 and 2002
    Report of Rapid Biodiversity Assessments at Cenwanglaoshan Nature Reserve, Northwest Guangxi, China, 1999 and 2002 Kadoorie Farm and Botanic Garden in collaboration with Guangxi Zhuang Autonomous Region Forestry Department Guangxi Forestry Survey and Planning Institute South China Institute of Botany South China Normal University Institute of Zoology, CAS March 2003 South China Forest Biodiversity Survey Report Series: No. 27 (Online Simplified Version) Report of Rapid Biodiversity Assessments at Cenwanglaoshan Nature Reserve, Northwest Guangxi, China, 1999 and 2002 Editors John R. Fellowes, Bosco P.L. Chan, Michael W.N. Lau, Ng Sai-Chit and Gloria L.P. Siu Contributors Kadoorie Farm and Botanic Garden: Gloria L.P. Siu (GS) Bosco P.L. Chan (BC) John R. Fellowes (JRF) Michael W.N. Lau (ML) Lee Kwok Shing (LKS) Ng Sai-Chit (NSC) Graham T. Reels (GTR) Roger C. Kendrick (RCK) Guangxi Zhuang Autonomous Region Forestry Department: Xu Zhihong (XZH) Pun Fulin (PFL) Xiao Ma (XM) Zhu Jindao (ZJD) Guangxi Forestry Survey and Planning Institute (Comprehensive Tan Wei Fu (TWF) Planning Branch): Huang Ziping (HZP) Guangxi Natural History Museum: Mo Yunming (MYM) Zhou Tianfu (ZTF) South China Institute of Botany: Chen Binghui (CBH) Huang Xiangxu (HXX) Wang Ruijiang (WRJ) South China Normal University: Li Zhenchang (LZC) Chen Xianglin (CXL) Institute of Zoology CAS (Beijing): Zhang Guoqing (ZGQ) Chen Deniu (CDN) Nanjing University: Chen Jianshou (CJS) Wang Songjie (WSJ) Xinyang Teachers’ College: Li Hongjing (LHJ) Voluntary specialist: Keith D.P. Wilson (KW) Background The present report details the findings of visits to Northwest Guangxi by members of Kadoorie Farm and Botanic Garden (KFBG) in Hong Kong and their colleagues, as part of KFBG's South China Biodiversity Conservation Programme.
    [Show full text]
  • GQ61 (2)Internet.Vp
    Geo log i cal Quar terly, 2017, 61 (2): 267–275 DOI: http://dx.doi.org/10.7306/gq.1344 Pol len morphotype edmundi and its sig nif i cance for palaeoclimate re con struc tions of the Neo gene Barbara S£ODKOWSKA1, * and Maria ZIEMBIÑSKA-TWORZYD£O2 1 Polish Geo log i cal In sti tute – Na tional Re search In sti tute, Rakowiecka 4, 00-975 Warszawa, Po land 2 Uni ver sity of War saw, Fac ulty of Ge ol ogy, ¯wirki i Wigury 93, 02-089 Warszawa, Po land S³odkowska, B., Ziembiñska-Tworzyd³o, M., 2017. Pol len morphotype edmundi and its sig nif i cance for palaeoclimate re con - struc tions of the Neo gene. Geo log i cal Quar terly, 61 (2): 267–275, doi: 10.7306/gq.1344 Pol len anal y sis has re vealed the pres ence of the edmundi morphotype in the Paleogene and Neo gene sed i ments of Cen tral Eu rope. Its spe cies, charac ter ized by dif fer ent morphologies, are mainly as signed to two fam i lies of ex tant plants – the Araliaceae and the Mastixiaceae – and com pared to pol len of some gen era (Diplopanax and Mastixia from the Mastixiaceae, and Aralia, Fatsia and Schefflera from the Araliaceae). The most sig nif i cant ge nus for palaeoclimate re con struc tions is the ex tant Diplopanax, whose seeds, sim i larly as the seeds of other rep re sen ta tives of the Mastixiaceae, ap pear in the warm phases of the Neo gene.
    [Show full text]
  • A Survey of Tricolpate (Eudicot) Phylogenetic Relationships1
    American Journal of Botany 91(10): 1627±1644. 2004. A SURVEY OF TRICOLPATE (EUDICOT) PHYLOGENETIC RELATIONSHIPS1 WALTER S. JUDD2,4 AND RICHARD G. OLMSTEAD3 2Department of Botany, University of Florida, Gainesville, Florida 32611 USA; and 3Department of Biology, University of Washington, Seattle, Washington 98195 USA The phylogenetic structure of the tricolpate clade (or eudicots) is presented through a survey of their major subclades, each of which is brie¯y characterized. The tricolpate clade was ®rst recognized in 1989 and has received extensive phylogenetic study. Its major subclades, recognized at ordinal and familial ranks, are now apparent. Ordinal and many other suprafamilial clades are brie¯y diag- nosed, i.e., the putative phenotypic synapomorphies for each major clade of tricolpates are listed, and the support for the monophyly of each clade is assessed, mainly through citation of the pertinent molecular phylogenetic literature. The classi®cation of the Angiosperm Phylogeny Group (APG II) expresses the current state of our knowledge of phylogenetic relationships among tricolpates, and many of the major tricolpate clades can be diagnosed morphologically. Key words: angiosperms; eudicots; tricolpates. Angiosperms traditionally have been divided into two pri- 1992a; Chase et al., 1993; Doyle et al., 1994; Soltis et al., mary groups based on the presence of a single cotyledon 1997, 2000, 2003; KaÈllersjoÈ et al., 1998; Nandi et al., 1998; (monocotyledons, monocots) or two cotyledons (dicotyledons, Hoot et al., 1999; Savolainen et al., 2000a, b; Hilu et al., 2003; dicots). A series of additional diagnostic traits made this di- Zanis et al., 2003; Kim et al., 2004). This clade was ®rst called vision useful and has accounted for the long recognition of the tricolpates (Donoghue and Doyle, 1989), but the name these groups in ¯owering plant classi®cations.
    [Show full text]
  • Relationships Within Cornales and Circumscription of Cornaceae—Matk and Rbcl Sequence Data and Effects of Outgroups and Long Branches
    MOLECULAR PHYLOGENETICS AND EVOLUTION Molecular Phylogenetics and Evolution 24 (2002) 35–57 www.academicpress.com Relationships within Cornales and circumscription of Cornaceae—matK and rbcL sequence data and effects of outgroups and long branches (Jenny) Qiu-Yun Xiang,a,* Michael L. Moody,b Douglas E. Soltis,c Chaun zhu Fan,a and Pamela S. Soltis d a Department of Botany, North Carolina State University, Raleigh, NC 27695-7612, USA b Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269-4236, USA c Department of Botany and the Genetics Institute, University of Florida, Gainesville, FL 32611-5826, USA d Florida Museum of Natural History and the Genetics Institute, University of Florida, Gainesville, FL 32611, USA Received 9 April 2001; received in revised form 1 March 2002 Abstract Phylogenetic relationships in Cornales were assessed using sequences rbcL and matK. Various combinations of outgroups were assessed for their suitability and the effects of long branches and outgroups on tree topology were examined using RASA 2.4 prior to conducting phylogenetic analyses. RASA identified several potentially problematic taxa having long branches in individual data sets that may have obscured phylogenetic signal, but when data sets were combined RASA no longer detected long branch problems. tRASA provides a more conservative measurement for phylogenetic signal than the PTP and skewness tests. The separate matK and rbcL sequence data sets were measured as the chloroplast DNA containing phylogenetic signal by RASA, but PTP and skewness tests suggested the reverse. Nonetheless, the matK and rbcL sequence data sets suggested relationships within Cornales largely congruent with those suggested by the combined matK–rbcL sequence data set that contains significant phylogenetic signal as measured by tRASA, PTP, and skewness tests.
    [Show full text]
  • In Vitro Culture of Camptotheca Acuminata (Decaisne) in Temporary Immersion System (TIS): Growth, Development and Production of Secondary Metabolites
    In vitro culture of Camptotheca acuminata (Decaisne) in Temporary Immersion System (TIS): Growth, development and production of secondary metabolites DISSERTATION A thesis submitted for the degree of Dr. rer.nat. (rerum naturalium) to the Biology Department, the Faculty of Mathematics, Informatics and Natural Sciences, University of Hamburg prepared by Yantree Devi Sankar-Thomas Republic of Guyana 2009 This thesis is dedicated to my husband and two children Table of Contents Page I. ABSTRACT I II. Zusammenfassung IV 1. INTRODUCTION 1 1.1. Aims of this Study 3 2. LITERATURE OVERVIEW 5 2.1. Camptotheca acuminata 5 2.1.1. Botany 5 2.1.2. Geographical Distribution 6 2.1.3. Ecology 7 2.2. Camptothecin (CPT) 8 2.2.1. Plant Secondary Metabolites 8 2.2.2. Camptothecin and its Pharmacological Effects 8 2.2.3. Semi-Synthetic Derivatives of Camptothecin 9 2.2.4. Biosynthetic Pathway of Camptothecin 10 2.3. Plant Micropropagation 12 2.3.1. Temporary Immersion System (TIS) 13 3. MATERIAL AND METHODS 17 3.1. Plant Material 17 3.1.1. Cuttings and Seeds 17 3.2. Plant Culture Media 18 3.3. Culture Vessels 19 3.3.1. Temporary Immersion System (TIS) Vessels 19 3.4. Sterilisation 20 3.4.1. Culture Apparatus, Glassware and Equipment 20 3.4.2. Nutrient Media and Substrates 21 3.4.3. Sterilisation of Plant Material 21 3.5. Growth Chambers 21 3.5.1. Illumination and Temperature 21 3.6. Plant Propagation Methods of C. acuminata 22 3.6.1. Ex Situ Seed Germination 22 3.6.2.
    [Show full text]
  • A Review of the Genus Curtisia (Curtisiaceae)
    Bothalia 39,1: 87-96 (2009) A review of the genus Curtisia (Curtisiaceae) E. YU. YEMBATUROVA* +, B-E. VAN WYK* • and P. M. TILNEY* Keywords: anatomy, Comaceae, Curtisiaceae, Curtisia dentata (Burm.f.) C.A.Sm., revision, southern Africa ABSTRACT A review of the monotypic southern African endemic genus Curtisia Aiton is presented. Detailed studies of the fruit and seed structure provided new evidence in support of a close relationship between the family Curtisiaceae and Comaceae. Comparisons with several other members of the Comales revealed carpological similarities to certain species of Comus s.I., sometimes treated as segregate genera Dendrobenthamia Hutch, and Benthamidia Spach. We also provide information on the history of the assegai tree, Curtisia dentata (Burm.f.) C.A.Sm. and its uses, as well as a formal taxonomic revision, including nomenclature, typification, detailed description and geographical distribution. INTRODUCTION (Feder & O’Brien 1968) was applied. Suitable sections were photographed. Fruits obtained from carpological Curtisia Aiton is a monotypic genus traditionally placed collections were rehydrated and then softened by means in the family Comaceae. It is of considerable interest of prolonged heating in Strassburger mixture (water, because of the many uses of its timber and bark—but no glycerol and 96 % ethyl alcohol in equal proportions), in recent reviews of the morphology, taxonomy or anatomy accordance with traditional anatomical procedures (Pro- are available. Recent cladistic and molecular systematic zina 1960) and then sectioned either by hand or sledge studies have revealed new evidence of relationships at fam­ microtome. Test-reactions to identify lignification (phloro- ily level (Murrell 1993; Xiang et al.
    [Show full text]
  • Mineralogy of Eocene Fossil Wood from the “Blue Forest” Locality, Southwestern Wyoming, United States
    Article Mineralogy of Eocene Fossil Wood from the “Blue Forest” Locality, Southwestern Wyoming, United States George E. Mustoe 1,*, Mike Viney 2 and Jim Mills 3 1 Geology Department, Western Washington University, Bellingham, WA 98225, USA 2 College of Natural Sciences Education and Outreach Center, Colorado State University, Fort Collins, CO 80523, USA; [email protected] 3 Mills Geological, 4520 Coyote Creek Lane, Creston, CA 93432, USA; [email protected] * Correspondence: [email protected] Received: 7 December 2018; Accepted: 7 January 2019; Published: 10 January 2019 Abstract: Central Wyoming, USA, was the site of ancient Lake Gosiute during the Early Eocene. Lake Gosiute was a large body of water surrounded by subtropical forest, the lake being part of a lacustrine complex that occupied the Green River Basin. Lake level rises episodically drowned the adjacent forests, causing standing trees and fallen branches to become growth sites for algae and cyanobacteria, which encased submerged wood with thick calcareous stromatolitic coatings. The subsequent regression resulted in a desiccation of the wood, causing volume reduction, radial fractures, and localized decay. The subsequent burial of the wood in silty sediment led to a silicification of the cellular tissue. Later, chalcedony was deposited in larger spaces, as well as in the interstitial areas of the calcareous coatings. The final stage of mineralization was the precipitation of crystalline calcite in spaces that had previously remained unmineralized. The result of this multi-stage mineralization is fossil wood with striking beauty and a complex geologic origin. Keywords: fossil wood; Blue Forest; Eden Valley; Lake Gosiute; chalcedony; quartz; calcite; stromatolite; Green River Basin; Wyoming 1.
    [Show full text]
  • ABSTRACT Fan, Chuanzhu. Molecular
    ABSTRACT Fan, Chuanzhu. Molecular Phylogeny and Evolution of Dogwoods. (Under the direction of Dr. Jenny Qiu-Yun Xiang) Dogwoods consist of morphologically diverse plants, and taxonomic circumscription and phylogenetic relationships of dogwoods have long been controversial. My dissertation study has two major goals: 1) elucidate phylogenetic relationships in the dogwoods group using nuclear DNA sequences, and 2) investigate the sequence evolution and its morphological link of the myc-like anthocyanin regulatory gene and explore the phylogenetic utility of the gene in dogwoods. Phylogenetic relationships within Cornus and Cornales (Cornus and related genera and families) were previously investigated using chloroplast DNA sequence data in several studies, but these still remained incompletely resolved. I used nuclear 26S rDNA sequences to further elucidate relationships within the group and to corroborate previously published phylogenetic hypotheses based on cpDNA and morphological data. Phylogenetic analyses of 26S rDNA sequence data (~3.4 kb) in combination with sequences of chloroplast genes rbcL and matK, suggest that the aquatic enigmatic genus, Hydrostachys from southern Africa, is sister to the remainder of Cornales among which Cornus and Alangium are sisters, nyssoids (Nyssa, Camptotheca, and Davidia) and mastixioids (Mastixia and Diplopanax) are sisters, and Hydrangeaceae and Loasaceae are sisters. These relationships, except the placement of Hydrostachys, are consistent with previous findings from analyses of matK-rbcL sequence data. Within Cornus, the dwarf dogwoods (subg. Arctocrania) are the sister of the big-bracted dogwoods (subg. Cynoxylon and subg. Syncarpea). This clade is, in turn, sister to the cornelian cherries (subg. Cornus and subg. Afrocrania). This large red-fruited clade is sister to a clade consisting of the blue- or white-fruited species (subg.
    [Show full text]
  • Cornus, Curtisia, Davidia, Diplopanax, Mastixia, and Nyssa Without Any Robust Lineages Among Them
    IAWA Journal, Vol. 19 (1),1998: 43-97 SYSTEMATIC WOOD ANATOMY OF CORNACEAE AND ALLIES by S. Noshiro l & P. Baas2 SUMMARY The wood anatomy of Comaceae, Alangiaceae, Garryaceae, and Nyssa­ ceae constituting the Comales in the sense of Cronquist (1981, 1988) is described in great detail and subjected to a cladistic analysis. A micro­ scopic identification key to the woods studied is given. The alliance in­ cludes seventeen genera, mostly of trees and shrubs, very rarely herbs. Although wood anatomically fairly homogeneous, variation exists in both qualitative and quantitative characters. Some of the latter show distinct latitudinal trends within individual genera, and character states have only been recognised taking their latitudinal dependencies into ac­ count. The character states ultimately recognised in these continuously varying quantitative characters coincide with intergeneric or intersec­ tional gaps. The cladistic analysis based on a datamatrix with twenty­ one characters (Table 3) and using Cereidiphyllum, Daphniphyllum, and Hamamelis as outgroups yielded a strict consensus tree with a quadrichotomy with two monophyletic clades, Hydrangea panieulata (a representative of the closely allied Hydrangeaceae) and Daph­ niphyllum (Fig. 81). One weakly supported clade includes Alangium, Camptotheea, Cornus, Curtisia, Davidia, Diplopanax, Mastixia, and Nyssa without any robust lineages among them. The other genera, Ara­ lidium, Aueuba, Corokia, Garrya, Griselinia, Helwingia, Melanophylla and Toricellia, constitute a second, well-supported clade. Two Hydran­ gea taxa included in the analysis nest in the second clade and a basal branching respectively. The wood anatomical diversity pattern thus supports a family concept of Comaceae including Cornus, Curtisia, Diplopanax, Mastixia, Alangiaceae, and Nyssaceae, and exclusion of the genera in the other clade.
    [Show full text]