Timber Trees of Suriname by LM Publishers

Timber Trees of Suriname An identification Guide

TIMBER Chequita R. Bhikhi TREES OF co-authors SURINAME

• IDENTIFICATION

Paul J.M. Maas

GUIDE •

Jifke Koek-Noorman Chequita R. Bhikhi Tinde R. van Andel Paul J.M. Maas editor Jifke Koek-Noorman Marion J. Jansen-Jacobs Tinde R. van Andel

Timber Trees of Suriname An identification Guide

Chequita R. Bhikhi co-authors Paul J.M. Maas Jifke Koek-Noorman Tinde R. van Andel editor Marion J. Jansen-Jacobs

Contents 4 Preface/ Voorwoord

Descriptive part

6 Introduction 10 Explanation of the descriptions

Anacardiaceae 24 Anacardium spruceanum 26 Loxopterygium sagotii 28 Tapirira guianensis

16 Illustrated glossary: plant parts

Apocynaceae 30 Aspidosperma album

20 Illustrated glossary: wood terms

Araliaceae 32 Schefflera morototoni Bignoniaceae 34 Handroanthus capitatus 36 Handroanthus serratifolius 40 Jacaranda copaia Boraginaceae 42 Cordia alliodora Burseraceae 44 Protium decandrum 46 Protium neglectum 48 Protium polybotryum 50 Tetragastris altissima 52 Tetragastris hostmannii 54 Trattinnickia burserifolia 56 Trattinnickia rhoifolia Caryocaraceae 58 Caryocar nuciferum Chrysobalanaceae 62 Couepia caryophylloides 64 Licania heteromorpha var. heteromorpha 66 Licania majuscula 68 Parinari campestris Clusiaceae 70 Garcinia benthamiana 72 Garcinia macrophylla 74 Garcinia madruno 78 Platonia insignis 80 Symphonia globulifera

Combretaceae 82 Buchenavia tetraphylla 84 Terminalia amazonia 86 Terminalia dichotoma 88 Terminalia guyanensis Fabaceae 90 Abarema jupunba 92 Andira inermis 94 Andira surinamensis 98 Bocoa prouacensis 100 Cedrelinga cateniformis 102 Copaifera guyanensis 104 Dicorynia guianensis 106 Diplotropis purpurea 110 Dipteryx odorata 112 Dipteryx punctata 114 Eperua falcata 116 Eperua grandiflora subsp. grandiflora 118 Eperua rubiginosa var. rubiginosa 120 Hydrochorea corymbosa 122 Hymenaea courbaril var. courbaril 128 Hymenolobium flavum 130 Inga alba 132 Martiodendron parviflorum 134 Mora excelsa 136 Ormosia coccinea 138 Parkia nitida 140 Parkia pendula 142 Peltogyne paniculata subsp. pubescens 142 Peltogyne paniculata subsp. paniculata 144 Peltogyne venosa subsp. densiflora 144 Peltogyne venosa subsp. venosa 146 Platymiscium ulei 148 Pseudopiptadenia suaveolens 154 Vatairea guianensis 156 Vataireopsis surinamensis 158 Vouacapoua americana Goupiaceae 160 Goupia glabra

Humiriaceae 164 Humiria balsamifera 166 Sacoglottis guianensis Lauraceae 168 Aniba panurensis 170 Aniba rosiodora 172 Licaria cannella 174 Ocotea glomerata 176 Ocotea guianensis 178 Ocotea percurrens 180 Rhodostemonodaphne grandis 182 Sextonia rubra Lecythidaceae 188 Bertholletia excelsa 190 Couratari guianensis 192 Eschweilera coriacea 194 Lecythis corrugata 196 Lecythis idatimon Loganiaceae 202 Antonia ovata Malvaceae 204 Lueheopsis rosea 2 06 Sterculia pruriens Meliaceae 208 Carapa guianensis 210 Carapa surinamensis 212 Cedrela odorata Moraceae 218 Bagassa guianensis 220 Brosimum guianense 222 Brosimum rubescens Myristicaceae 224 Virola michelii 226 Virola surinamensis Putranjivaceae 230 Drypetes variabilis Salicaceae 232 Laetia procera

Cedrelinga cateniformis - slash. Photo: C.R. Bhikhi

Sapotaceae 234 Manilkara bidentata subsp. bidentata 234 Manilkara bidentata subsp. surinamensis 236 Micropholis guyanensis 238 Pouteria cuspidata subsp. robusta 238 Pouteria cuspidata subsp. dura 2 40 Pradosia ptychandra

264 Glossary 270 Translation list English-Dutch of wood and botanical terms 275 Index of scientific names 277 Index of vernacular names 280 Notes 281 Literature

Simaroubaceae 2 42 Simarouba amara Vochysiaceae 2 48 Erisma uncinatum 250 Qualea albiflora 252 Qualea coerulea 254 Qualea dinizii 256 Qualea rosea 258 Vochysia guianensis 260 Vochysia tomentosa

Preface Alberta (“Bep”) M.W.Mennega

It has been some 50 years since Bomenboek voor Suriname, authored by three botanists from the Herbarium of Utrecht, the Netherlands, was first published. The Bomenboek described about 100 commercially important species of trees native to Suriname and included short descriptions of many related species in order to facilitate reliable identification. The leading author of the first edition was Jan C. Lindeman, an ecologist with a vast knowledge of the flora and vegetation of Suriname. JCL – to his colleagues – had collected and identified numerous plants from Suriname and was able to identify almost all the tree species. He was renowned for being able to recognise a taxon on leaf material alone. The second author, Alberta (“Bep”) M.W.Mennega, was the Herbarium’s official wood anatomist. When sterile plant material attached to a bit of wood was sent to the Herbarium, Bep would take her large pocketknife and slice a sliver from the top of the piece of wood and, using her hand lens, suggest the correct family or even genus name. At the time Bep and JCL were the two leading authorities on the flora of Suriname. The third contributor who deserves a mention is Wim H.A.Hekking. Wim was a plant taxonomist who had dedicated his career to the taxonomy of the tropical woody genera of the Violaceae. He was also a skilled botanical artist and his beautifully detailed drawings ensured the Bomenboek was totally user-friendly. The Bomenboek has been, and is still today, an invaluable aid to all those studying the Suriname flora and vegetation. As a young botanist in 1964-65, I undertook many vegetation studies in the forests of western Suriname. At that time the tropical rainforest appeared to me simply as a large tangle of unidentifiable tree trunks where seldom could either a flower or fruit be found. Fortunately for me, trained “boomkenners”, Rudolf Elburg and John Tawjoeran, were often on site calling out names such as “walaba, 15 cm diameter”, or “alata-oedoe, 18 cm diameter”. I would then immediately turn to the Bomenboek to check these indigenous names and only then understand that they had spotted Catostemma fragrans, or Minquartia guianensis. However all was not as simple as it seemed for when they called “pisi” or “guave” for example I learned all too quickly that these names could refer to any number of different species. In such cases a small leafy twig and a small piece of wood would be collected and sent to the Netherlands where the Utrecht Herbarium

staff would be able identify the species that had so confused me in the field. Since 1963 many people have relied heavily on the Bomenboek but for many years it has been out of print. Two years ago, however some 50 years after its first publication, the Alberta Mennega Stichting – a foundation that awards grants to young botanists to help with financing fieldwork in the tropical or publishing research relating to tropical botany celebrated its 25th anniversary. To mark this occasion the board of the Alberta Mennega Stichting decided to facilitate the publication of the second edition of the book to which Bep Mennega had contributed so much. Moreover the board decided that this new edition should appear in English rather than Dutch so that more people could benefit from the wealth of information it contained. Chequita Bhikhi, a Suriname biologist and expert on the flora of her homeland, was willing to undertake this daunting task and revise the book in close collaboration with her co-authors. The focus was once again on ca 100 species of commercially important trees native to Suriname but in this edition it was decided that information relating to the commercially less important tree species would be omitted. This allowed more space to expand the botanical descriptions of those tree species that were retained and include important information pertaining to bark features for nearly all of the species. All descriptions in this newly revised edition are richly illustrated with photographs and botanical drawings. I speak on behalf of the board members of the Alberta Mennega Stichting when I say I am confident that this book will continue to be an invaluable resource for botanists, taxonomists and all those interested in the woody flora of both Suriname and the wider South American tropics. May, 2016 P.J.M.Maas www.alberta-mennega-stichting.nl

Het is nu ruim 50 jaar geleden (1963) dat de eerste versie van het “Bomenboek voor Suriname” verscheen. Het boek werd geschreven door drie bekende botanici van het Utrechtse Herbarium en beschreef ongeveer 100 boomsoorten, die voor de handel belangrijk hout leveren. Als vergelijking waren korte beschrijvingen van verwante, maar commercieel minder belangrijke soorten opgenomen. De eerste auteur was Jan C. Lindeman, een Utrechtse plantenecoloog met een enorme kennis van de flora en vegtetatie van Suriname. JCL, zoals zijn Utrechtse collega’s hem noemden, had zelf tijdens zijn vegetatiekundig onderzoek in Suriname veel plantenmateriaal verzameld en kende vrijwel alle boomsoorten. Hij herkende het materiaal, zelfs als er alleen maar blad aanzat, dus zonder bloemen of vruchten. De tweede auteur was Alberta (“Bep”) M.W. Mennega, houtanatome. Wanneer er in het Utrechtse Herbarium plantenmateriaal binnenkwam van een boom uit Suriname, zonder bloemen of vruchten maar wel met een houtmonster, dan gebruikte zij haar grote zakmes, sneed er een plakje af, keek met haar loupe en wist dan meestal meteen tot welke familie of zelfs geslacht de boom behoorde. Zij en JCL waren in die tijd in Utrecht dé twee grote vraagbakens op het gebied van de Surinaamse flora. De derde persoon die heel belangrijk heeft bijgedragen aan het Bomenboek was Wim H.A. Hekking. Wim was een echte plantentaxonoom, zijn hele carrière werkend aan de tropische houtige vertegenwoordigers van familie Violaceae. Daarnaast was hij ook botanisch tekenaar. Door zijn fraaie illustraties was het Bomenboek zeer gebruikersvriendelijk. Het Bomenboek was en is een enorme hulp voor onderzoekers die de Surinaamse flora en vegetatie bestuderen. Zo was ik zelf (in 1964/65) in Suriname voor vegetatiekundig onderzoek aan de bossen in het Westen van dat land en had daar, als beginnende plantentaxonoom, een enorme steun aan het Bomenboek. Het tropische regenbos was toen voor mij nog één grote wirwar van onbekende boomstammen, waaraan je maar zelden een bloem of vrucht zag. Gelukkig had ik hulp van de zeer kundige boomkenners Rudolf Elburg en John Tawjoeran, die me bij de bosinventarisaties dagelijks toeriepen “walaba, 15 cm diameter”, “alata-oedoe, 18 cm diameter” enz, enz. Bij het noemen van die namen keek ik onmiddellijk in het Bomenboek (voor 5 Surinaamse guldens in Paramaribo bij Varenkamp gekocht) en wist dan dat het om

Catostemma fragrans en Minquartia guianensis ging. Wanneer zij echter “pisi” of “gujave” riepen bleek uit het Bomenboek dat er vele soorten waren die pisi en gujave konden heten en dan werd er voor controle onmiddellijk een bebladerde tak, plus een stukje hout voor Bep Mennega, verzameld. Daarna kon in het Utrechtse Herbarium worden geverifieerd om welke soorten het ging. Heel veel onderzoekers hebben het Bomenboek sinds 1963 gebruikt, helaas is het nu al vele jaren uitverkocht. Twee jaar geleden, ca. 50 jaar nadat het Bomenboek werd gepubliceerd, bestond de Alberta Mennega Stichting 25 jaar. Deze stichting helpt jonge botanici speciale onderdelen van hun onderzoek te financieren, zoals veldwerk, of bijzondere publicaties, alles met betrekking tot tropische botanie. Het leek het bestuur van de stichting een goed idee om het boek, waaraan Bep Mennega zo’n essentiële bijdrage had geleverd, een “opvolger” te geven. De Surinaamse biologe Chequita Bhikhi, die tijdens haar studie al veel onderzoek had verricht in Suriname en de flora ervan goed kende, bleek bereid om deze klus op zich te nemen, in nauwe samenwerking met de co-auteurs. In korte tijd is de informatie over zo’n 100 belangrijke boomsoorten herzien en uitgebreid met veel botanische details. Voor vrijwel alle soorten konden ook de bastkenmerken worden vermeld. Besloten werd om het boek ditmaal niet in het Nederlands, maar in het Engels te laten verschijnen, waardoor een grotere gebruikersgroep bereikt kan worden. De korte beschrijvingen van verwante soorten zijn nu weggelaten, waardoor meer ruimte kwam voor illustraties. Zo zijn alle beschrijvingen nu voorzien van botanische tekeningen, kleurenfoto’s, en macroscopische houtfoto’s. Namens de Alberta Mennega Stichting hoop ik, dat het boek een belangrijke bron van informatie zal zijn voor allen met belangstelling voor de flora van tropisch Amerika, Suriname in het bijzonder. Mei 2016 P.J.M. Maas www.alberta-mennega-stichting.nl

timber trees of suriname

Voorwoord

Introduction Suriname, covering an area of 16.4 million hectares and forming part of the Guiana Shield basement complex, is situated on the Northern coast of South America. Approximately 94% of the total land surface is covered with forest with about 27% (4.5 million hectares) reserved for timber production purposes (sbb 2014; fao 2015). Tree species are harvested from this concession area, which is situated in the “Forest belt” in northern Suriname (Figure 1). In 2003, this area totaled 103 concessions covering a land area of ca. 1.6 million hectares. Conventional prospecting, often referred to as “tree spotting”, involves identification of potential trees in a concession and is often carried out prior to harvesting (Forest Department 2001). Identification of trees is undertaken by experienced spotters trained in tree identification based on bark and slash characteristics. However differences between the species within one family may not be recognized by bark and slash morphology only. Leaves, flowers and fruits are of utmost importance when identifying any plant species. This book focuses on the identification of Surinamese trees based on field, vegetative, floristic and wood characteristics. It includes botanical descriptions, wood descriptions, illustrations and photographs of 100 Surinamese

commercial timber tree species, potential timber tree species and tree species protected by Surinamese forest law (S.B. 2000 no. 208). This book is therefore intended for anyone interested in learning about and identifying Surinamese timber trees such as timber companies, forest management organizations, students and conservationists. Whilst this book focuses on the trees of Suriname, many of the species treated herein can also be found in adjacent Guyana and French Guiana, whilst others have a much wider Neotropical distribution still. For more than 50 years the ‘Bomenboek voor Suriname’ (Lindeman & Mennega 1963) has been the only book focusing on the identification of commercial and non-commercial trees of Suriname based on wood and vegetative characteristics. Although the “Bomenboek voor Suriname” is outdated, up until now it has been the best Figure 1. Map showing the Forest belt, where timber harvesting takes place in Suriname (from “Stichting voor Bosbeheer en Bostoezicht” or the Foundation for Forest Management and Production Control (sbb) Suriname (2015), reproduced with permission)

7 timber trees of suriname

Qualea dinizii - trunk base with buttresses and flaky bark. Photo: P. & S. Teunissen

identification guide for timber trees of Suriname. Since the “Bomenboek voor Suriname” is no longer in publication it has become important for a new timber trees identification book to be published and for that book to carry updated information regarding timber tree species, vegetative descriptions and wood anatomical descriptions. This new book focuses on 100 commercial and potential timber trees including eight tree species protected by Surinamese forest law (S.B. 2000 no. 208). The selection of species is based on ‘Jonkers (1987) checklist of important commercial species’ published in Annex 1 of Werger (2011) supplemented by current (2007-2013) Surinam Forest Service (sbb) production and export data of wood and wood products of tree species with a high commercial value. Furthermore the species list has been compared with the list of commercial timber species (Group a) and species prohibited from felling (Group c) according to the Surinamese forest law (S.B. 2000 no. 208). In this book we provide information on the bark, twigs, leaves, flowers, fruits and wood of the 100 timber species covered to facilitate easy identification in the field. The great majority of the descriptions are accompanied by scientific illustrations and colour photographs of plant parts. Although the information provided in this book should be sufficient for the identification of trees in the field, it is recommended that dried plant material housed in the local herbarium is also consulted. Not only will this provide a complete picture of the plant, but certain features such as hair covering, hair types, or differences between leathery and papery leaves can more easily be observed on dried material.

Qualea rosea - trunk base with buttresses. Photo: PC.R. Bhikhi

Acknowledgements This book is an initiative of- and funded by the Alberta Mennega Stichting. I would like to express my gratitude to the board of the Alberta Mennega Stichting who gave me the opportunity to work for the last 16 months on this wonderful project. I would like to thank the Naturalis Biodiversity Center (my former employer) for facilitating support during the project. Home and work commitments meant that I had to undertake much travelling to and from the Herbarium of Naturalis Biodiversity Center in Leiden to gather literature and meet my supervisors, therefore I am very grateful for a grant from the Stichting het Van Eeden-Fonds that helped cover part of the travel expenses. I would like to thank Stichting Kronendak, Berch van Heemstede Stichting, Hugo de Vries Fonds, Stichting het Van Eeden-fonds, Naturalis Biodiversity Center and Tropenbos International Suriname all for the financial support towards the publication of this book.

I am gratefully indebted to Scott Mori for reviewing the descriptions of the Lecythidaceae species and for allowing us to use many photographs and illustrations; Douglas Daly for his generous support in reviewing the descriptions of the Burseraceae species and for granting permission to use a number of photographs as well; Sir Ghillean Prance for reviewing the descriptions of the Chrysobalanaceae species; Terry Pennington for reviewing the descriptions of the Sapotaceae species and granting permission to use the illustration of Pradosia ptychandra; and Henk van der Werff for reviewing and commenting on the descriptions of the Lauraceae species. Thanks are also due to Naturalis Biodiversity Center Leiden for allowing us to use the illustrations by Hendrik Rypkema and the late Wim Hekking. I am also grateful to Bobbi Angell for allowing us to use a number of her illustrations in this book, and Esmée Winkel for her hard work in preparing the last illustrations within a tight time frame. I would also like to thank André van Proosdij (Naturalis Biodiversity Center), Pieter Schriks (Walburg Pers), the Royal Botanic Gardens Kew, David H. Boshier (University of Oxford), and the Missouri Botanical Garden Press for granting permission to include illustrations that were originally published in van Proosdij (2001), Boshier and Lamb (1997), Stace (2009), Marcano-Berti (1997) and Woodson et al. (1967).

< Lecythis corrugata - bark. Photo: P. & S. Teunissen

Special thanks go to Pieter and Sila Teunissen, Joelaika Ramdas-Behari, Guno Marjanom and Fabian Diran who are responsible for most of the photographs. I would also like to thank Carol Gracie for letting us use her copyrighted photographs that were originally published in Mori et al. (2002). I thank Sofie Ruysschaert, Steven Paton and Rolando Pérez (Smithsonian Tropical Research Institute), Domingos Cardoso (Instituto de Biologia Universidade Federal da Bahia (ufba)) and Robin Foster for sharing many of their photographs as well. Special thanks are also due to Sarah Crabbe (SBB Suriname) for permitting the use of the ‘Forest belt’ map in the Introduction and for providing us with valuable wood production data that helped us make the rigorous selection of timber trees for this book. Thank you Frits van Troon for your help with the determinations of many trees during a regional workshop at Tonka Island that enabled me to make additional bark descriptions and photographs for a number of trees that were lacking. Thanks also go to Olaf Bánki for his support, enthusiasm and endless discussions about trees and forest types that ensured a lasting interest in this project; Imogen Poole is gratefully acknowledged for correcting and translating the introductory parts of this book; Berte van Heuven (Naturalis Biodiversity Center) for preparing wood slides for the book; Leo Junikka for valuable comments and explanations concerning the terminology used to make accurate bark descriptions; Hiltje Maas-van de Kamer and Pieter Baas for their support. In addition, I thank Maureen Playfair (celos Suriname), Rudi van Kanten (Tropenbos Suriname) and the National Herbarium of Suriname all for their support and interest in this project. Chequita R. Bhikhi

timber trees of suriname

This book would not have been possible without the guidance and continuous support of my advisors and co-authors, Paul Maas and Jifke Koek-Noorman, for making plant and wood descriptions in particular as well as for all their valuable comments throughout the project: thank you both very much. I am grateful to Tinde van Andel (co-author), as project coordinator, for her overall support especially concerning the vernacular names used in this book, and for acquiring the funding for this project; Marion Jansen-Jacobs, editor, for her general advice and support in finalising the original drawings; Lubbert Westra who is responsible for all the wood images; Sylvia Mota de Oliveira for providing me with valuable literature needed to make the descriptions and for support concerning additional illustrations and permissions.

Explanation of the descriptions Scientific names and synonyms Family names of all species described follow the APG III system of the Angiosperm Phylogeny Group (2009). Scientific species names were verified with ‘The Plant List’ - a working list of all known plant species (version 1.1, 2013) from the Royal Botanic Gardens Kew, the Missouri Botanical Garden and “Tropicos.org” from the Missouri Botanical Garden (2015). According to these lists some species have many synonyms, but we include the most up to date ones mentioned in recent publications. For example, Handroanthus serratifolius (Vahl) S.O.Grose has 23 synonyms, we only mention Tabebuia serratifolia (Vahl) G.Nicholson that has been used extensively in publications.

Vernacular names The most commonly used vernacular names in Suriname are cited in alphabetical order with the abbreviation given in parentheses behind. Vernacular names have been taken mostly from van ´t Klooster et al. (2003), Lindeman & Mennega (1963) and the “Flora of Suriname” and “Flora of the Guianas” series. Vernacular names are mentioned in eight different languages. The abbreviations of these languages are: Ar = Arawak, Au = Aucan, Ca = Caribbean, Du = Dutch, Pa = Paramaccan, Sa = Saramaccan, SD = SurinamDutch, Sr = Sranangtongo.

Botanical description Each botanical description begins with a field description of the tree followed by a detailed description of leaves, inflorescence, flowers, fruits and seeds of the plant. Field description Field description includes the mean height of the tree, the trunk diameter at breast height (DBH) of the mature tree, trunk shape and the presence or absence of buttresses. Detailed description of bark is provided where ever possible and is taken from Polak (1992), Grenand & Loubry (2001), Mennega (1948), Lindeman & Mennega (1963) and supplemented with information from herbarium material, colour photographs of slashes and some field observations. The terminology used for bark descriptions follows the book “Survey for English macroscopic bark terminology” by Junikka (1994). Information is included regarding the bark surface, the outer bark (layer between the inner bark and the bark surface) and the inner bark (living part of the bark). The description of the bark surface includes the colour, texture, patterns (fissuring, scaling) and

external markings (such as lenticels, hoops). The outer bark is described in terms of texture, structure and thickness. The colour of the outer bark is only mentioned when it differs from the colour of the bark surface. The inner bark is described in terms of colour, thickness, structure, texture, and exudation (colour, thickness and smell). The field description ends with a general description of the twigs, which includes the shape in cross section, colour, indument and thickness of the twigs. A ‘thin’ twig is ca. 2 mm in diameter whereas a ‘thick’ twig is nearer ca. 4 mm. Plant description Leaves, inflorescences, flowers and fruits are described in detail but are limited to characters that can be seen with the naked eye or hand lens. All descriptions are taken from Floras and manuscripts. Those consulted most often were “Flora of the Guianas”, “Flora of Suriname”, “Flora Neotropica”, “Flora of the Venezuelan Guayana”, “Guide to the vascular plants of Central French Guiana” (Mori et al. 1997, 2002), “Bomenboek voor Suriname” (Lindeman & Mennega 1963), “Fruits of the Guianan Flora” (van Roosmalen 1985), and various publications with family or species treatments. Additional information and measurements were taken from herbarium material and labels. The descriptions of the plant families Burseraceae, Chrysobalanaceae, Lauraceae, Lecythidaceae, and Sapotaceae were reviewed by scientists specialising in these families. At the end of this book, a glossary including botanical and wood terms used in this book is given. In an Illustrated Glossary (p. 16-22), schematical drawings of some plant parts and of wood features can be found. The Illustrated Glossary is based on the “Bomenboek voor Suriname” (Lindeman & Mennega 1963), and “Arnoldo’s Zakflora” (van Proosdij 2001). The terminology for leaf shapes follows that outlined in the “Terminology of simple symmetrical plane shapes chart 1a” (Systematics Association Committee for Descriptive Biological Terminology 1962). Commonly used leaf shape terminology are elliptic, oblong, rhombic, ovate, obovate and triangular with ratios of 2:1 and 3:2 between leaf length and width. Other commonly used leaf shape terminology are narrowly elliptic, narrowly oblong, narrowly rhombic, narrowly ovate, narrowly obovate and narrowly triangular with ratios of 6:1 and 3:1, linear with a ratio of 12:1, circular with a ratio of 1:1 and deltate with ratios of 1:1 and 5:6.

11 timber trees of suriname

Carapa surinamensis - flowering twig. Photo: P. & S. Teunissen

Ecology and distribution

Photographs

The section ‘Ecology and distribution’ includes information regarding forest types, species occurrence, distribution in the three Guianas and the Neotropics, and finally types of seed dispersal. The description of forest types is generally based on Lindeman & Mennega (1963), Mori et al. (1997, 2002), Polak (1992), Japing (1957), Japing & Japing (1960, 1961), van Roosmalen (1985) and in some cases data taken from labels of herbarium material. Occurrence and seed dispersal data have mostly been taken from van Roosmalen (1985). Distribution data generally come from “GBIF portal” (http://www.gbif.org/species), “Tropicos” (www.tropicos.org), “Discover life” (http:// www.discoverlife.org/), “List of species of the Brazilian flora” (http://floradobrasil.jbrj.gov.br/), the “Checklist of the plants of the Guiana Shield” (Funk et al. 2007) and the Floras mentioned above.

Most of the plant images used in this book were derived from tree specimens stored in the herbarium of Naturalis Biodiversity Center in Leiden, the Netherlands, and in the National Herbarium of Suriname (BBS). Collection numbers of these specimens are written between parentheses and included in the caption for each image. Most images can be found below the descriptions. Other images are to be found on photo pages with a reference to them in the description of the species. Each wood image is reproduced from a 35 mm slide that was taken at 5 x magnification, and is provided with a scale bar of 500 μm. Special characters, such as oil cells in rays, are identified on some images.

Notes In the notes section a summary of the main differences between subspecies and varieties of the species dealt with is given. The subspecies and varieties are provided between parentheses and follow the species name.

Illustrations Most illustrations by H. Rypkema are drawn on a scale of 1:1 enabling comparisons between different species. In many illustrations the original size of the plant part is given. The botanical artists responsible for the majority of the illustrations are Hendrik Rypkema, Wim Hekking, Esmée Winkel and Bobbi Angell. Most illustrations by H. Rypkema are reprinted from Polak (1992), those by W.H.A. Hekking are reprinted from Lindeman & Mennega (1963) with permission from Naturalis Biodiversity Center, and

Symphonia globulifera - flowering twigs. Photo: C.R. Bhikhi

illustrations by Bobbi Angell are reprinted, with permission, from Mori et al. (2002). The illustrations by Esmée Winkel were drawn specifically for this book. We attempted to add illustrations of all species described but unfortunately we could not provide high quality and complete illustrations for all species - some illustrations are out dated and taken from old manuscripts and some, especially those of W.H.A. Hekking, are not complete.

Wood description The wood descriptions are based on information from the literature. Important sources are: “Bomenboek voor Suriname” (Lindeman & Mennega 1963), “Major Timber Trees of Guyana: Wood Anatomy” (Miller & Détienne 2001), “Major Timber Trees of Guyana: Timber Characteristics and Utilization” (Gérard, Miller & ter Welle 1996), the “Insidewood database (2004onwards)” from the North Carolina State University (NCSU) published on the internet (http://insidewood. lib.ncsu.edu/search), “Suriname Timbers” (Mennega 1948), “Suriname Timber Species: Characteristics and utilization” (Comvalius 2001), “Onderzoek naar de belangrijkste mechanische en fysische eigenschappen van 41 Surinaamse houtsoorten” (Japing 1957), “Houthandboek Surinaamse Houtsoorten” (Japing & Japing 1960), “Handleiding voor hout, in het bijzonder Surinaamse houtsoorten” (Japing & Japing 1961) and “Hout van Leguminosae uit Suriname” (ReindersGouwentak & Rijsdijk 1968).

Photographs of cross sections are given for all species. Some features are explained in the chapter “Illustrated Glossary”. For further details and pictures of the features as well as the terminology used in this book, we refer the reader to the “IAWA list of microscopic features for hardwood identification” (IAWA Committee 1989, also available on internet e.g. https://www.researchgate.net/ publication/294088872_IAWA_List_of_Microcopie_ Features_for_Hardwood_Identification). The wood descriptions focus on features than can be seen either with the naked eye or with a handlens with a magnification of 10 x or 20 x. At this magnification it is possible to see cross, radial and tangential sections of the wood in great detail (Mennega 1948) (Figure 19). The cross section follows the plane perpendicular to the axis of the trunk, the radial section is the vertical plane through the axis towards the bark, and the tangential plane runs parallel to the bark of the tree. A sharp (pocket) knife, good light conditions and a measuring slide such as the “loepmicrometer” in the “Bomenboek voor Suriname”, which can be downloaded from www.lmpublishers/xxx are required. The “loepmicrometer” is divided in square millimetres, 2 series of parallel lines between 0.5 mm (500 µm) and 0.2 mm, and a series of parallel lines of 30 µm, 50 µm, 100 µm, and 200 µm. It is important that the three sections are prepared properly in order to ensue clear observations with a hand lens. The cross- and tangential sections of the wood sample

General properties Colour The colour of the wood is observed from the cross section. Differences in colour between sapwood and heartwood, as well as the discolouration by the influence of light or aging, are mentioned in addition to other characters such as flame marks, streaks or blotches. Texture The texture is observed on the tangential section. Texture is an indication for the fineness of the wood and is influenced by the width and the number of vessels. Wide vessels may form conspicuous longitudinal grooves, which may result in a coarser texture. If the grooves are fine and close together, the surface will be smooth and the texture fine. Three classes are distinguished: coarse, medium, fine. Grain The grain can be observed on the tangential and radial section of the wood. Grain is closely related to the texture of the wood and describes the course and direction of the vessels and fibres with respect to the axis of the tree - this is often important as it affects woodworking techniques (e.g. against the grain). Descriptions and grain types include: straight, spiral and interlocked. Straight grain: axial wood elements run parallel to the axis of the tree. Spiral grain: axial wood elements make an angle to the axis of the tree. Interlocked grain: the orientation of the axial wood elements with respect to the axis of the tree changes regularly resulting in an irregular surface to radial splits. Wood with interlocked grain is common in tropical wood and sometimes characteristic in certain families such as the Vochysiaceae where it appears in almost every tree.

seasonal tree growth takes place annually. Tropical trees tend to have more than one growing season each year depending on the rainy and dry seasons and may form more than one complete or incomplete growth ring in a year (Lindeman & Mennega 1963; Mennega 1948). Although we indicate whether growth rings are distinct, indistinct, or absent, differences may in fact be gradual. Distinct growth ring boundaries are marked by differences in the number and size of vessels, in the distribution of parenchyma and in the colour and density of the fibre tissue. Often indistinct growth rings are marked by slightly gradual structural changes at poorly defined or barely visible boundaries. Vessels Vessels are vertical tubes, formed by vessel elements, serving as pipelines within the trunk and responsible for the transport of sap within a tree. Vessel elements are the largest type of wood cells and can generally be seen with the naked eye. In cross section vessels can be identified as square, round or oval openings. In the descriptions the following vessel characteristics are recorded: porosity, vessel arrangement, vessel grouping, perforation plates, tangential diameter of vessels, number of vessels per square millimetre, tyloses and deposits in vessels. Porosity Hardwoods are divided into three main categories according to the number, size and arrangement of their vessels: ring-porous, semi-ring-porous and diffuse-porous. In diffuse-porous woods - by far the most common type - vessels and vessel groups are evenly distributed and vessel diameters are more or less uniformly distributed throughout the wood. In ring-porous wood the large earlywood vessels form a clearly defined ring with an abrupt transition at the junction with the latewood. In semi-ring-porous woods the large earlywood vessels gradually change to narrower latewood vessels.

Wood structure

Vessel groupings Vessels may be solitary, or occur in radial multiples or in clusters (Figure 21). Exclusively solitary: more than 90% of vessels appear to have no contact with another vessel. Radial multiples: files formed by two to many adjacent vessels. Clusters: groups of 3 to many radially and tangentially adjacent vessels. The most common vessel grouping is a combination of solitary vessels and radial multiples of two to four pores.

Growth rings Growth rings in tropical trees are not as distinct and regular as they are in trees of temperate zones where

Vessel arrangement Instead of being evenly scattered, vessels and vessel

Odour and taste These features are only mentioned when present. Odour can be a valuable feature in certain families. The original odour of dry wood may return on making a new cut.

timber trees of suriname

should be cut with a sharp knife, whereas the radial section should be fractured. It is also important for the wood sample to be thoroughly dry, although sometimes a drop of water may be helpful to see details on the cross and tangential sections.

groups may be arranged in straight or wavy tangential bands, or in radial, diagonal or zig-zag patterns (Figure 21). Perforation plates Perforation plates are the openings in the end walls of adjacent vessel elements that allow fluids to pass from one vessel element to another along the length of the vessel. Perforation plates are difficult to recognize with a hand lens and are best viewed in radial sections. In tropical woods simple perforation plates with a single opening are most common. Multiple perforation plates can be scalariform (Figure 22) or reticulate. Reticulate perforation plates have a netlike appearance and are common in Bignoniaceae. When they are simple, the type of perforation plate is not mentioned in the description. Vessel diameter The vessel diameter is defined as the width of the opening in the tangential direction as seen in cross section. Vessels should be measured at the widest part of the opening, excluding the wall. The most frequent range of width is given as well as minimum and maximum widths. At least 25 vessels should be measured. Smaller vessels should be excluded, except for when larger and smaller vessels are evenly distributed or when there is a striking difference in width between early wood and late wood vessels. The latter case occurs, for example, in Cedrela odorata where two measurements are given in the descriptions, one for the early wood and one for the late wood. Vessel frequency As with vessel diameter the number of vessels is variable within a species and should be measured with the square millimeter division of the measuring slide. Each vessel should be counted as one individual and thus radial multiples of 3 vessels should be counted as 3 vessels. Counts from at least 5-10 fields of 1 sq. mm spread over many growth rings should be taken . For larger vessels, larger fields or whole fields of view at low magnification should be counted (IAWA Committee 1989; Lindeman and Mennega 1963). In the descriptions the frequency range is given alongside the minimum and maximum values. Tyloses and deposits Tyloses are bubble-like outgrowths from adjacent ray or axial parenchyma cells that grow into open vessels (Figure 22D). As with gums or deposits of different origin and colour, they can partially or completely block the vessels in the heartwood. In the description

the tylose walls (thin, thick or sclerotic) are also recorded. Rays Rays consist of parenchyma cells that run from the centre of the axis out to the bark. They serve to transport nutrients between the cambium, sapwood and pith (Meier 2008-2015). The structure of rays is observed on the radial split section of wood. In cross section rays can be seen as lines running from the centre to the bark, and in tangential section as narrow lines or lenses (Figure 19). Three main types of ray cells are distinguished: procumbent, square, and upright (Figure 23B). Rays can be composed of one-, or a combination of two or three different cell types. Often rays are composed of multiseriate central parts composed of procumbent cells with uniseriate margins of 1-4 or more rows of upright and/or square cells. In some species procumbent, square and upright cells are mixed throughout the ray. Rays are very important for the identification of wood by virtue of their great variety in structure, number and size. Ray width and height The width and height of rays is observed in the tangential section and is given as number of cells and/or in micrometers (Îźm) (Figure 23C, D). In tropical woods rays are seldom more than 1 mm high. The descriptions frequently include the estimated minimum and maximum height. Number of rays per millimeter The number of rays is best counted from the cross section. The average of at least 10 measurements should be taken. In the descriptions the minimum and maximum number per mm are given. Parenchyma Axial parenchyma can be seen in cross section (Figure 19) and is identified by its lighter colour when compared with the surrounding fibre tissue. Parenchyma patterns are one of the most important features for the identification of wood (Mennega 1948) because of the wide variety that can be found. Parenchyma may be found as diffuse cells or very short to long lines of cells unassociated with the vessels (apotracheal parenchyma). Parenchyma patterns clearly associated with the vessels (paratracheal parenchyma) can vary from narrow, incomplete rings around the vessel to complete (vasicentric) rings or wide rings with confluent extensions. Vasicentric parenchyma with lateral extensions may be lozenge-aliform with diamond-shaped outline, winged-aliform with narrow, elongated extensions and confluent with coalescing

Parinari campestris - part of inflorescence with flowers. Photo: C.R. Bhikhi, CRB 136

extensions forming irregular bands. Both apotracheal and paratracheal parenchyma can form parenchyma bands that can be short to long, narrow to wide, straight to wavy, few to numerous, or may form a regular network with the rays (Figure 20). Marginal parenchyma can be found at the margin of growth rings as a more or less continuous layer of variable width. Storied structure Storied structure is formed when wood cells are arranged in horizontal series or tiers (Figure 23A). When formed by rays, tiers are recognisable as fine horizontal striations or ripple marks visible on the tangential section with the naked eye. Although it is a useful feature for identifying wood, variability in storied structure can be found within a species and between samples (IAWA Committee 1989). Special features Apart from the features listed above, additional wood features that may help to recognize a genus or family include included phloem, oil cells, intercellular canals, latex or tanniniferous tubes, and mineral inclusions. Included phloem Small amounts of phloem or bark tissue included within the wood because of abnormal growth of the cambium, may appear on the cross section as concentric tangential bands or scattered islands, e.g. Antonia ovata, Erisma uncinatum (IAWA Committee 1989) (Figure 24A).

Intercellular canals and tubes In many families secondary plant products such as latex, gums, resins, are found in tubes or in intercellular canals. Both intercellular canals and tubes may extend either radially within a ray or axially between the ground tissue. The difference between intercellular canals and intercellular tubes can only be observed under the microscope. Intercellular canals are surrounded by epithelial cells that may produce gums or resin. Tubes are formed by a series of cells and may contain latex or, rarely, tannins. Radial tubes and radial canals often appear as dark lines in the rays in radial section and as small dark spots in tangential section (Figure 24C, D). Axial canals are often enclosed by parenchyma and arranged in short to long tangential bands or lines, or distributed in either a solitary fashion or in small clusters (Mennega 1948; IAWA Committee 1989) (Figure 24E). They are observed in cross section and are recognizable by their irregular shape, arrangement and colour. When they are formed in response to injury their occurrence is variable. Mineral inclusions The following mineral inclusions are mentioned in the descriptions: prismatic crystals, styloids, elongate crystals and silica bodies. Prismatic crystals of calcium oxalate are the most common crystal type and can be located in rays, axial parenchyma and among fibres (IAWA Committee 1989); they are birefringent and best observed under polarized light. Styloids are large crystals at least four times as long as they are broad with either pointed or squared ends. Elongate crystals are two to four times as long as broad with pointed ends. Silica bodies are small, globose or irregular particles occurring in rays, parenchyma or fibres. Consisting of silicon dioxide these are not birefringent and are best observed in radial sections with a light microscope.

timber trees of suriname

Oil cells Oil cells are cells filled with oil of different colours varying from yellowish through to red, occasionally bright orange, sometimes empty, often enlarged. They are characteristic of certain families, for example Lauraceae. They can be associated with rays, parenchyma or may be present among the fibres. They often occur as enlarged cells among marginal cells of rays in tangential and radial section (Figure 24B).

Illustrated glossary: plant parts

Figure 2. Simple leaves

Figure 3. Compound leaves

Figure 4. Leaf arrangement

17 timber trees of suriname

Figure 5. Leaf shape (reprinted from van Proosdij (2001), with permission of A.S. J. van Proosdij and Walburg Pers, Zutphen (2015))

Figure 6. Leaf apex

Figure 7. Leaf base

Figure 8. Leaf margin

Figure 9. Leaf venation pattern

Figure 10. Primary vein cross section

Figure 11. Petiole cross section

Figure 12. Stipules

Figure 13. Hair types

from van Proosdij (2001), with permission of A.S. J. van Proosdij and Walburg Pers, Zutphen (2015))

Figure 15. Flower shape (reprinted from van Proosdij (2001), with permission of A.S. J. van Proosdij and Walburg Pers, Zutphen (2015))

Figure 16. Longitudinal section of flowers

Figure 17. Sexual reproduction

Figure 18. Longitudinal section of fruit

timber trees of suriname

Figure 14. Inflorescence (reprinted

Illustrated glossary: wood terms

Figure 19. Dicotyledonous wood sample showing tangential, radial and cross section 1. Vessels with simple perforation plates 2. Fibre tissue 3. Rays composed of procumbent body ray cells, with one row of upright cells at the margins 4. Banded parenchyma

Figure 20. Parenchyma patterns in cross section of wood Apotracheal parenchyma: A. diffuse or diffuse-in-aggregates, e.g. Sacoglottis guianensis. B. banded, in narrow continuous tangential bands or lines, evenly spaced but narrower than the rays, e.g. Micropholis guyanensis. C. banded, reticulate, in continuous tangential bands or lines of about the same width as the rays, evenly spaced, and forming a network with them, e.g. Bertholletia excelsa. Paratracheal parenchyma: D. vasicentric, e.g. Sextonia rubra. E. aliform, winged-aliform and aliform-confluent, e.g. Brosimum rubescens. F. lozenge-aliform and confluentbanded, e.g. Parkia pendula.

Vessel groupings: A. Vessels exclusively solitary, e.g. Aspidosperma album. B. Vessels solitary and in short radial multiples (of mostly 2 or 3 cells), e.g. Qualea rosea. C. Vessels solitary, in radial multiples of 2-5(-7) and small clusters, e.g. Ormosia coccinea. D. Vessels solitary, and in short and long radial multiples (of 2 or more cells). Vessel arrangement: D. Vessels in radial pattern, e.g. Manilkara bidentata. E. Vessels in diagonal pattern, e.g. Sextonia rubra. F. Vessels in tangential bands indicating a growth zone.

Figure 22. Vessel elements and tyloses A-C. Perforation plates (pe) of vessel elements. (A, B) simple, (C) scalariform with 6 bars (b) D. Tyloses (t): outgrowths from adjacent ray cells into the vessel.

timber trees of suriname

Figure 21. Vessel groupings and arrangement in cross section of wood

Figure 23. Rays A. Storied structure of rays, with all rays storied (tangential section), e.g. Simarouba amara. B-E. Cellular composition of rays. (B) Multiseriate ray composed of procumbent (p) body ray cells, with one to three rows of upright (u) and/or square (s) marginal cells. (C) Multiseriate ray surrounded by larger cells or sheath cells (sh), as viewed on the tangential section, e.g. Cordia alliodora. (D) Uniseriate ray composed of upright or square cells. (E) Multiseriate portions composed of procumbent body ray cells, linked to long uniseriate extensions of square cells or upright cells.

Figure 24. Special features A. Included phloem, diffuse: isolated phloem strands scattered in the fibre tissue, e.g. Antonia ovata (cross section) B. Oil cells associated with ray margins (between upright and/or square marginal cells), e.g. Ocotea glomerata (radial section). C-E. Resin canals. C-D. Radial canal present in multiseriate ray, e.g. Tapirira guianensis. E. Axial resin canals (c) included in parenchyma band, e.g. Lecythis corrugata (cross section).

Species are arranged in alphabetical order by family names, then by genus names.

Trunk of Dipteryx odorata. Photo: C.R. Bhikhi

timber trees of suriname

Descriptive part

Anacardium spruceanum Benth. ex Engl. ANACARDIACEAE Vernacular names

Ecology and distribution

boskasju (Sr), busi-kasju (Sr), busi kasyun (Pa).

In non-inundated rain forest and forest next to rivers. Uncommon. The 3 Guianas, Bolivia (Pando), Amazonian Brazil (Amapá, Amazonas, Maranhão, Pará, Rondônia), and Venezuela (Amazonas, Bolívar). Zoochorous.

Botanical description Tree to 20-35 m tall, trunk 60-180 cm in diam., fairly straight, cylindrical, and slightly swollen at base; bark pale grey, smooth, with scattered lenticels, inner bark reddish brown, resin red; twigs with prominent cordate leaf scars and erect, golden coloured hairs. Leaves alternate, aggregated toward branch tips; petiole 15-40 mm long, flattened, glabrous; blade papery, obovate, 9-27 x 3.5-11 cm, glabrous on both sides, base acute, sometimes obtuse, margin entire, flat, apex rounded or obtuse, sometimes slightly emarginate, truncate, acuminate or mucronate; primary vein impressed above, secondary veins in 13-16 pairs, slightly raised above. Inflorescences terminal, broadly pyramidal with dense tufts of flowers, 5-20 x 3-20 cm, covered with soft, brownish hairs toward distal branches; bracts leafy; peduncle 0.5-4 cm long; pedicels 1.5-3 mm long, sparsely covered with soft hairs. Flowers bisexual and staminate, sparsely pubescent. Bisexual flowers: 5-merous; sepals 5, narrowly ovate to angular-ovate, 3-7 mm long; petals 5, pink to dark purple, more or less linear, 6-12 mm long, apex reflexed; stamens 8-10, basally fused into a staminal tube of 1-5 mm long, 1 stamen much longer, 5-9 mm long; ovary superior, 1-locular, subglobose, ovule 1, basal, style 1, terminal. Staminate flowers: similar size as bisexual flower. Fruit a drupe, black when mature, subreniform, 1-3 x 2-4 cm, attached to fleshy, edible pedicel or hypocarp; hypocarp white, red, or yellow, pyriform, 1-2.5 x 2-3 cm, with strong resinous smell.

Fruits with fleshy, edible pedicel. Photo: C.R. Bhikhi

Wood description Sapwood greyish white with more or less pinkish tinge, slightly distinct from the yellowish to pale or russet brown heartwood. Grain slightly interlocked and irregular, texture medium to coarse. Growth rings indistinct or absent. Vessels diffuse-porous, solitary and in radial multiples of 2-3, up to 100-210 μm wide, 1-5 per sq. mm. Tyloses thin-walled, common. Rays 7-11 per mm, exclusively uniseriate, up to ca. 1 mm (22-26 cells) high; body ray cells procumbent with (1-)2-4 rows of upright and/or square marginal cells. Silica bodies present. Parenchyma paratracheal, in narrow vasicentric rings, aliform or lozenge-aliform, occasionally confluent.

Wood sample (Maas LBB 10735, Uw 11193)