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‘establishment strategy’ = recruitment, regeneration Seminar Series GreenMondays 05 September 2011 IB-UPD Seedling establishment in tropical rain forests: understanding the evolutionary strategies Edwino S. Fernando PhD College of Forestry and Natural Resources THE UNIVERSITY OF THE PHILIPPINES - LOS BAÑOS Page . 1 Page . 2 Hazards of young tree In the forest, seeds and seedlings tend to be . Establishment strategies seedlings in the forest: • eaten or defoliated by / mechanisms for animals (herbivory), seedling survival • buried deep under 1 Germination stem and leaf litter, rate • washed away by rain, 2 Seedling • trampled by animals architecture or falling tree branches, (initial seedling morphology / functional • infected or killed by morphology) pathogenic microbes, or 3 Initial seedling • starved by lack of size HopeaPage . 3acuminata Merr. light, water, or nutrients. ShoreaPage . astylosa4 Foxw. DIPTEROCARPACEAE DIPTEROCARPACEAE (Ng 1978, Leishman et al. 2000, Westoby et al. 2002) Germination rate Germination rates, viability, and relative weight of Dipterocarpaceae fruits from the Makiling Botanic Gardens, Philippines (Lapitan 1996) Rapid germination - Germination rate Normal length of Number of seeds Species total germination of all (days) viability (weeks) per 100 gm viable seeds within 12 Anisoptera aurea 7-11 2-3 57 weeks of seed fall; c. Anisoptera thurifera 14-21 4-8 167-174 65% of species sampled Dipterocarpus 30 2-5 6-7 grandiflorus • standard based on seed germination of Hopea acuminata 7 1 - dipterocarps Hopea malibato 20 2-8 962-1000 • all Malayan dipterocarp Hopea plagata 1-6 3 148-151 seeds germinated within Parashorea 14-21 2 70-74 12 weeks (Ng 1978) malaanonan Shorea almon 1-16 2 32-33 all Philippine dipterocarp • Shorea contorta 5-12 1-2 14-15 seeds germinated within Memecylon edule Roxb. Shorea polysperma 3-10 3 108-110 Page . 5 MELASTOMATACEAE 1-4 weeks (Lapitan 1996) Page . 6 1 Species with rapid Rapid germination - within the main stream of evolution in tropical rain forests germination rate Most vulnerable to logging High concentration of seedlings necessary because the forest to ensure the survival of a few individuals floor does not hold a reservoir of their seeds. Survivors likely to be found in areas Can simplify nursery of highest seedling density; management; rapid fruits/m2 rate of turnover more food than ensures that nursery predators can space and labor are consume not tied up for long periods. Page . 7 Page .Ng8 (1978) Rapid germination Rapid germination - within the main stream of Hopea malibato: evolution in tropical rain forests fruit density on the The ‘Janzen-Connell ground and dispersed effects’ High concentration of seedlings necessary distance from mother disproportionately high tree to ensure the survival of a few individuals mortality of seedlings near parent trees or adults because of host- Survivors likely to specific predators, be found in areas at herbivores, or pathogens the edge of the distance- or density- dependent predation dispersal area where seedling fruits/m2 fruits/m2 density is lowest; less likely to be attractive to predators; less competition among seedlings Page . 9 HopeaPageSuzakimalibato . 10 et Foxwal. 1986. Suzaki et al. (1986) Ng (1978) DIPTEROCARPACEAE Rapid germination Parashorea malaanonan: Rapid germination Shorea contorta: seedling seedling density and density and dispersed dispersed distance from 2 mother tree distance from mother tree Shorea contorta Vidal DIPTEROCARPACEAE Seedlings/m m 2 Seedlings/m m Parashorea Suzaki et al. (1986) Page . 11 Suzaki et al. (1986) Page . 12 malaanonan (Blanco) Merr. DIPTEROCARPACEAE 2 Germination rate Parashorea malaanonan: seedling / sapling size class Prolonged germination – m distribution beneath mother tree germination after at least 12 weeks of dormancy (delayed germination) or germination that begins before 12 weeks, but do not 15 year old achieve total germination within 12 weeks (intermediate germination); c. 8 year old 35% of species sampled Seedlings produced few at a time over an extended period 6 year old Predator pressure and 3 year old competition among seedlings 1 year old reduced Long-lasting reservoir of seeds on the forest floor; can be moved by animals or cm Parashorea rain, effective dispersal area enlarged PageSuzaki . 13et al. (1986) malaanonan (Blanco) Merr. Page . 14 Gomphandra mappioides Valeton DIPTEROCARPACEAE Ng (1978) STEMONURACEAE Tree seedling architecture Tree seedling architecture (Initial seedling morphology) Nomenclature of tree seedling parts (initial seedling morphology in four types of germination) Elaeocarpus monocera ELAEOCARPACEAE 1 root phanerocotyl -ar epigeal, Anisoptera aurea with DIPTEROCARPACEAE epicotyl 2 collet or root collar foliaceous or reserve cryptocotyl- ar hypogeal, cotyledons cotyledons 3 hypocotyl (stem axis below the cotyledons) with reserve cotyledons 4 cotyledons or seed leaves hypocotyl hypocotyl (may be ‘bulky’ or ‘reserve’ type, or hypocotyl developed, undeveloped, foliaceous or leaf-like) cotyledons cotyledons exposed unexposed 5 epicotyl (stem axis above the cotyledons) 6 cataphylls or scale leaves 7 eophylls (first fully expanded leaves) Swietenia macrophylla MELIACEAE 8 stem epigeal hypogeal Ng (1978) PageFernando . 15 (1980) Page . 16 Tree seedling architecture (initial seedling morphology in four types of germination) epicotyl phanerocotyl cryptocotyl- -ar hypogeal, cotyledons ar epigeal, with reserve with reserve cotyledons cotyledons hypocotyl hypocotyl undeveloped, hypocotyl developed, cotyledons cotyledons exposed unexposed Relative percentage of initial seedling morphological types in semi-hypogeal „durian‟ type PageNg . 17 (1978) PageMalayan . 18 forest trees (Ng 1978) 3 Relative percentage of initial seedling morphological (‘durian’ type ) D 10% CER types in Mexican forest trees (Ibarra-Manriquez et al. 2001) cryptocotylar Erythrina folkersii Pseudolmedia oxyphyllaria epigeal with Omphalea oleifera LEGUMINOSAE MORACEAE EUPHORBIACEAE C 3% reserve storage A 80% cotyledons B 7% 2.4% CHR (hypogeal) PER cryptocotylar hypogeal with Cojoba arborea CHR LEGUMINOSAE reserve storage CER cotyledons 31.4% Psychotria faxlucens PEF (epigeal) RUBIACEAE phanerocotylar PEF (semi- epigeal with PHR hypogeal) (epigeal) foliaceous PHR PER cotyledons phanerocotylar phanerocotylar hypogeal with Relative percentage of initial seedling morphological types in 49.5% epigeal with reserve reserve cotyledons cotyledons PageMakiling . 19 forest trees (Fernando 1980) Page . 20 7.2% 9.5% Seedling classification after Garwood (1996) Epigeal type of Epigeal type of initial seedling morphology initial seedling morphology allows the cotyledon to function as photosynthetic organs; seedlings can become independent of food reserves at an early stage common in species with small seeds (<3 mm along axis, excluding wings) the most common type in the tropical rain forests; also characteristic of pioneer species (e.g. Ficus, Trema) Terminalia foetidisssima Griff. Page . 21 Antidesma bunius (L.) Spreng. Page . 22 Celtis luzonica Warb. COMBRETACEAE De Vogel (1980) PHYLLANTHACEAE Ng (1978) ULMACEAE Fernando (1980) Epigeal type of Epigeal type of initial seedling morphology Ng (1978) Fernando (1980) initial seedling morphology Some species with epigeal germination have not exploited the photosynthetic advantage. In Diospyros, the cotyledons emerge and are shed off immediately without turning green; the first two leaves are developed Pittosporum ferrugineum W.T.Aiton PITTOSPORACEAE precociously, as if to compensate for the early loss Bischofia javanica Blume STAPHYLEACEAE Ng (1978) Page . 23 Diospyros blancoi A.DC. of the cotyledons. Page . 24 EBENACEAE 4 Epigeal type of Epigeal type of initial seedling morphology initial seedling morphology Canarium hirsutum Willd. BURSERACEAE Anisoptera aurea Foxw. DIPTEROCARPACEAE Shorea contorta Vidal De Vogel (1980) DIPTEROCARPACEAE Gomphandra mappioides Valeton Page . 25 Page . 26 Fernando (1980) STEMONURACEAE Epigeal type of Hypogeal type of initial seedling morphology initial seedling morphology • The resting of the seed body on the ground favors the development of bulky storage tissue; often the endosperm rather than the cotyledons. • Usually produce scale leaves (cataphylls), as if Photo: L.L. Co to emphasize the Shorea palosapis Merr. abundance of stored DIPTEROCARPACEAE Shorea malibato Foxw. Page . 27 Pagefood . 28 reserves. Atuna racemosa Raf. DIPTEROCARPACEAE Ng (1978) CHRYSOBALANACEAE Ng (1978) Hypogeal type of Hypogeal type initial seedling morphology of initial seedling morphology Pseudolmedia oxyphyllaria Don.Sm. MORACEAE cataphyll Lithocarpus sp. FAGACEAE Lithocarpus korthalsii Ng (1978) FAGACEAE Swietenia macrophylla King MELIACEAE De Vogel (1980) Fernando (1980) Scorodocarpus borneensis (Baill.) Becc. Barringtonia macrostachya (Jack) Kurz Page . 29 Page . 30 OLACACEAE LECYTHIDACEAE 5 Germination ‘adjacent-ligular’ Hypogeal type Germination Hypogeal type of initial seedling morphology ‘remote-ligular’, of initial seedling morphology Corner (1966) eophyll ‘remote-tubular’ soil level seed seed cataphyll soil or litter level hyperphyll ligule (coleoptile) May help to get radicle Orania longisquasma the seedling into ARECACAEAE the ground Sabal umbraculifera ARECACAEAE http://www.flickr.com/photos/mora Orania paraguanensis through the leaf beza79/2229424609/sizes/l/in/pho ARECACAEAE tostream/ litter Corner (1966) Areca costulata Page . 31 Page . 32 Areca costulata ARECACAEAE ARECACAEAE Semi-hypogeal type of
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    LARGE TREES, SUPERTREES AND THE GRASS PHYLOGENY Thesis submitted to the University of Dublin, Trinity College for the Degree of Doctor of Philosophy (Ph.D.) by Nicolas Salamin Department of Botany University of Dublin, Trinity College 2002 Research conducted under the supervision of Dr. Trevor R. Hodkinson Department of Botany, University of Dublin, Trinity College Dr. Vincent Savolainen Jodrell Laboratory, Molecular Systematics Section, Royal Botanic Gardens, Kew, London DECLARATION I thereby certify that this thesis has not been submitted as an exercise for a degree at any other University. This thesis contains research based on my own work, except where otherwise stated. I grant full permission to the Library of Trinity College to lend or copy this thesis upon request. SIGNED: ACKNOWLEDGMENTS I wish to thank Trevor Hodkinson and Vincent Savolainen for all the encouragement they gave me during the last three years. They provided very useful advice on scientific papers, presentation lectures and all aspects of the supervision of this thesis. It has been a great experience to work in Ireland, and I am especially grateful to Trevor for the warm welcome and all the help he gave me, at work or outside work, since the beginning of this Ph.D. in the Botany Department. I will always remember his patience and kindness to me at this time. I am also grateful to Vincent for his help and warm welcome during the different periods of time I stayed in London, but especially for all he did for me since my B.Sc. at the University of Lausanne. I wish also to thank Prof.
  • Dictionary of Cultivated Plants and Their Regions of Diversity Second Edition Revised Of: A.C

    Dictionary of Cultivated Plants and Their Regions of Diversity Second Edition Revised Of: A.C

    Dictionary of cultivated plants and their regions of diversity Second edition revised of: A.C. Zeven and P.M. Zhukovsky, 1975, Dictionary of cultivated plants and their centres of diversity 'N -'\:K 1~ Li Dictionary of cultivated plants and their regions of diversity Excluding most ornamentals, forest trees and lower plants A.C. Zeven andJ.M.J, de Wet K pudoc Centre for Agricultural Publishing and Documentation Wageningen - 1982 ~T—^/-/- /+<>?- •/ CIP-GEGEVENS Zeven, A.C. Dictionary ofcultivate d plants andthei rregion so f diversity: excluding mostornamentals ,fores t treesan d lowerplant s/ A.C .Zeve n andJ.M.J ,d eWet .- Wageninge n : Pudoc. -11 1 Herz,uitg . van:Dictionar y of cultivatedplant s andthei r centreso fdiversit y /A.C .Zeve n andP.M . Zhukovsky, 1975.- Me t index,lit .opg . ISBN 90-220-0785-5 SISO63 2UD C63 3 Trefw.:plantenteelt . ISBN 90-220-0785-5 ©Centre forAgricultura l Publishing and Documentation, Wageningen,1982 . Nopar t of thisboo k mayb e reproduced andpublishe d in any form,b y print, photoprint,microfil m or any othermean swithou t written permission from thepublisher . Contents Preface 7 History of thewor k 8 Origins of agriculture anddomesticatio n ofplant s Cradles of agriculture and regions of diversity 21 1 Chinese-Japanese Region 32 2 Indochinese-IndonesianRegio n 48 3 Australian Region 65 4 Hindustani Region 70 5 Central AsianRegio n 81 6 NearEaster n Region 87 7 Mediterranean Region 103 8 African Region 121 9 European-Siberian Region 148 10 South American Region 164 11 CentralAmerica n andMexica n Region 185 12 NorthAmerica n Region 199 Specieswithou t an identified region 207 References 209 Indexo fbotanica l names 228 Preface The aimo f thiswor k ist ogiv e thereade r quick reference toth e regionso f diversity ofcultivate d plants.Fo r important crops,region so fdiversit y of related wild species areals opresented .Wil d species areofte nusefu l sources of genes to improve thevalu eo fcrops .