Stratification of a tropical forest as seen in dispersal types

331 Pages·1986·25.212 MB·English

by  Ingrid Roth (auth.)

Stratification of a tropical forest as seen in dispersal types
Tasks for vegetation science 17
Series Editors
HELMUT LIETH HAROLD A. MOONEY
University of Osnabriick, F.R. G. Stanford University, Stanford, Calif" U.S.A.
I Box, E.O. Macroclimate and Plant Forms. An introduction to predictive modelling in phytogeography.
ISBN 90 6193 941 0
2 Navin Sen, D. & Singh Rajpurohit, K. Contributions to the Ecology of Halophytes. ISBN 90 6193 942 9
3 Ross, J. The Radiation Regime and Architecture of Plant Stands. ISBN 906193607 1
4 Margaris, N.S. & Mooney, H.A. (eds) Components ofP roductivity ofM editerranean - Climate, Regions.
ISBN 90 6193 944 5
5 Miiller, M.L Selected Climatic Data for a Global Set of Standard Stations for Vegetation Science.
ISBN 90 6193 945 3
6 Roth, I. Stratification in Tropical Forests as Seen in Leaf Structure. ISBN 90 6193 946 1
7 Steubing, L. & Jager, H.J. Monitoring of Air Pollutants by Plants: Methods and Problems.
ISBN 90 6193 947 X
8 Teas, H.J. Biology and Ecology of Mangroves. ISBN 90 61939488
9 Teas, H.J. Physiology and Management of Mangroves. ISBN 90 61939496
10 Feoli, E., Lagonegro, M. & Orl6ci, L. Information Analysis of Vegetation Data. ISBN 90 6193950 X
II Sestak, Z. (ed) Photosynthesis during Leaf Development. ISBN 90 6193951 8
12 Medina, E., Mooney, H.A. & Vazquez-Yanes, C. (eds) Physiological Ecology ofP lants oft he Wet Tropics.
ISBN 90 6193 952 6
13 Margaris, N.S., Arianoustou-Faraggitaki, M. & Oechel, W.C. (eds) Being Alive on Land.
ISBN 90 6193 953 4
14 Hall, D.O., Myers, N. & Margaris, N .S. (eds) Economics ofE cosystems Management. ISBN 90 61935059
15 Estrada, A. & Fleming, T.H. (eds) Frugivores and Seed Dispersal. ISBN 90 6193543 1
16 Dell, B., Hopkins, A.J.M. & Lamont, B.B. (eds) Resilience in Mediterranean-type Ecosystems.
ISBN 90 6193 5792
17 Roth, I. Stratification of a Tropical Forest as Seen in Dispersal Types. ISBN 906193613 6
Stratification
of a tropical forest
as seen in dispersal types
by
INGRID ROTH
1987 DR W. JUNK PUBLISHERS
a member of the KLUWER ACADEMIC PUBLISHERS GROUP
DORDRECHT I BOSTON I LANCASTER
Distributors
jor the United States and Canada: Kluwer Academic Publishers, P.O. Box 358,
Accord Station, Hingham, MA 02018-0358, USA
for the UK and Ireland: Kluwer Academic Publishers, MTP Press Limited, Falcon
House, Queen Square, Lancaster LAI lRN, UK
jor all other countries: Kluwer Academic Publishers Group, Distribution Center,
P.O. Box 322, 3300 AH Dordrecht, The Netherlands
Library of Congress Cataloging in Publication Data
Roth, Ingrid.
Stratification of a tropical forest as in
dispersal types.
(Tasks for vegetation science ; 17)
Bibliography: p.
Includes indexes.
1. Rain forest ecology. 2. Tropical plants-
Ecology. 3. Seeds--Tropics--Dispersal. 4. Fruit-
Tropics--Dispersal. 1. Title. II. Series.
QK938.F6R675 1986 581.5'2642 86-10489
ISBN-13, 978-94-010-8639-4 e-ISBN-13, 978-94-009-4826-6
DOI, 10.1007/978-94-009-4826-6
Copyright
© 1987 by Dr W. Junk Publishers, Dordrecht.
Softcover reprint of the hardcover 1st edition 1987
All rights reserved. No part of this publication may be reproduced, stored in a
retrieval system, or transmitted in any form or by any means, mechanical,
photocopying, recording, or otherwise, without the prior written permission of
the publishers,
Dr W. Junk Publishers, P.O. Box 163, 3300 AD Dordrecht, The Netherlands.
Die Schopfung des Lebendigen ist
auf eine Entwicklung hin geplant,
sonst hatte der T od keinen Sinn.
Dedicated to my beloved mother
PREFACE
Our knowledge of the structure and dispersal of "Plants, seeds and currents" in the Westindies
tropical fruits and seeds is very limited up to the and Azores. Van der Pijl with his extensive know
present day, though richness of species and variety ledge of tropical plants offers a great selection of
of forms is overwhelming in the tropical forests. detailed information on the subject" Principles of
Morphology of tropical fruits and seeds has always dispersal in higher plants" (1972, and earlier pa
attracted the curiosity of botanists from many pers). The author who has earned most merits in
countries and information may be obtained from the field of seed and fruit predation, chemical
books and publications concerned with taxonomy. defenses of plants, and animal-plant interactions is
Ulbrich's "Biologie der Friichte und Samen" Janzen. He and his collaborators have thrown new
(1928) gives examples of tropical fruits and seeds light on this subject. Nonetheless, a large unknown
and their dispersal methods. The two volumes by field still remains ahead of us, especially regarding
Van der Roosmalen (1977) dealing with the de a detailed knowledge of fruit and seed dispersal of
scription of tropical plants were of utmost value to tropical plant species.
me, as the area considered, Surinam, is close to The great opportunity for my own studies was
Venezuelan Guiana and because both regions have the fruit and seed collection of Dr. Bernard Rollet
many species in common. However, the inner to whom I am very much obliged for placing the
structure of tropical fruits and seeds is less known. material at my disposal. He made the collection
Some new examples of the structure and develop during his stay in Venezuelan Guiana when he
ment of tropical fruits and seeds have been pre directed the tree inventory project in this area. My
sented in my book "Fruits of Angiosperms" thanks go also to Dr. P. R. Mohana Rao who
(1977). Comer's "Seeds of Dicotyledons" (1976) started the morphological studies together with me
offers many detailed structural peculiarities of in Venezuela, but unfortunately was compelled to
seeds in drawings. Unfortunately, the least infor return to India soon after his arrival for health
mation is available regarding detailed descriptions reasons. Finally, I have to thank my former stu
offruit and seed dispersal of tropical plant species. dent and collaborator, Licenciado Ricardo Castro
In his bible "Plant dispersal throughout the Penso, for the preparation of the slides used for
world" Ridley (1930) gives a general survey over the work reported in this volume.
the most important facts. Guppy (1917) deals with
CONTENTS
PREFACE VII
1. SOURCE OF THE MATERIAL AND SAMPLE COLLECTION
2. INTRODUCTION 3
3. GENERAL PART 5
Fruit and seed types
Factors influencing the presence of plants
Seasonality of flowering and fruiting
Dispersal methods
Hydrochory (rain wash, floating fruits and seeds, submerged transport of diaspores in the water, dispersal by the
sea) -Anemochory -Autochory and explosively dispersed seeds -Dispersal by fish (ichthyochory) -Dispersal by
reptiles (saurochory) -Dispersal by birds (ornithochory) -Dispersal by mammals (mammaliochory except
chiropterochory) -Dispersal by bats (chiropterochory) -Dispersal by ants (myrmecochory)
Protection against predation
Mechanical protection -Chemical protection -Nutritional aspects -Escape in space and time
Coevolution of plants and animals
Germination and establishment
Seed size -Latency and viability -Germination -Vivipary
4. RESULTS 49
Description of the families studied concerning fruit and seed outer and inner structure, dispersal methods, fruiting
time, height of the tree, number of individuals, and special habitat
Acanthaceae -Anacardiaceae -Annonaceae -Apocynaceae -Araliaceae -Bignoniaccae -Bixaceae -Bombacaceae
-Boraginaceae -Burseraceae -Capparidaceae -Caryocaraceae -Celastraceae -Cochlospermaceae -Combretaceae
-Dichapetalaceae -Ebenaceae -Elaeocarpaceae -Euphorbiaceae -Aacourtiaceae -Guttiferae -Hernandiaceae -
Hippocrateaceae -Humiriaceae -Lacistemaceae -Lauraceae -Lecythidaceae -Leguminosae (Mimosaceae,
Caesalpiniaceae, Papilionaceae) -Linaceae -Malpighiaceae -Melastomaceae -Meliaceae -Moraceae -Myristicaceae
-Myrtaceae -Nyctaginaceae -Ochnaceae -Olacaceae -Opiliaceae -Palmae -Polygonaceae -Quiinaceae -
Rhamnaceae -Rhizophoraceae -Rosaceae -Rubiaceae -Rutaceae -Sabiaceae -Sapindaceae -Sapotaceae -
Simaroubaceae -Sterculiaceae -Tiliaceae -Verbenaceae -Violaceae -Vochysiaceae -Unidentified species without
family
Number of species and individuals studied and their height categories
Number of species and individuals studied and their dispersal methods
Plant families and their dispersal methods related to the height categories
Important characteristics of certain families, genera or species and their number of individuals
Fruiting times and dispersal mechanisms
Anemochory -Autochory -Auto-zoochory -Zoochory
Fruiting times at different heights (A-a-aa) of the forest
Families, dispersal types and fruiting times
Geographical distribution, habitats and fruiting times
Animal life and forest layering (Ground animals, tree animals, aquatic animals)
Seasonality of animal life in the forest
Dispersal methods in dry and humid regions
Forest layering and dispersal strategies
Proportion of fleshy to dry fruits in the different strata
Monochory and polychory
Species diversity and series of species
Refuge theory
Number of individuals and height categories in homogeneous groups
Evolutionary considerations
Forest stratification and the evolution of the heightening forest
Comparison of bark, leaf and fruit structure in the fumilies studied
Usefulness of tropical fruits and seeds
x
FINAL CONCLUSIONS 180
BIBLIOGRAPHY 191
INDEX OF VERNACULAR PLANT AND ANIMAL NAMES 313
INDEX OF SCIENTIFIC PLANT AND ANIMAL NAMES 317
GENERAL INDEX 323
CHAPTER I
SOURCE OF THE MATERIAL AND SAMPLE COLLECTION
The material investigated mainly comes from the inundated soils or permanently inundated river
humid tropical forest in Venezuelan Guiana where banks, rocky sites, etc., which may well be recog
an inventory was initiated by the Ministry of nized by their floristic composition. In its western
"Agricultura y Cria" (MAC) of Venezuela to part, the forest is in contact with a belt of semide
gether with the Food and Agriculture Organization ciduous and deciduous forests, bordering in their
(FAO) and the UNO. In this inventory all arbor turn, savannas. Although the tropical humid forest
eous species of this area were statistically covered. is called "evergreen", Rollet (1964) was able to
The total area studied for the inventory amounted count about 60 species which drop their leaves,
to about 67 km in lenght and approximately 25 m mainly in April. However, the defoliation is some
in width, covering in total about ISS,S ha. A clear what ephemeral and irregular, even considering
ing of the above mentioned length and width was one and the same species so that the forest as a
cut through the forest to collect trees, shrubs, and whole preserves its evergreen aspect. Flowering
partly also lianas. In total, 67.777 individual trees and fructification are not very clearly limited to
(and shrubs) with a diameter of not less than 10 certain periods of the year, but according to Rollet
cm at breast height were included. a certain proportion of the species flowers and
The tropical humid forest from which leaf sam fruits with more intensity four times a year, in
ples were collected corresponds to a dense ever February-June-August-December and in Februa
green rain forest which Beard would call" season ry-May-August-December, respectively. For more
al" to indicate the incidence of distinct dry sea details of the humid tropical forest in the State of
sons. Geographically it is situated on the shield of Bolivar, Venezuela, and its floristic composition,
Venezuelan Guiana at the so-called massif of Ima see Rollet (1964, 1969a, b), from which publica
taca (south of the Orinoco river and east of the tions the above data are taken.
junction with the Caroni river). This extreme nor Most of the species were identified by Dr. Julian
thern part of the Guianan shield has an undulated Steyermark, Instituto Botanico, Caracas. Those
ground relief which fluctuates between ISO m and samples which could not be identified in Caracas
550 m in altitude. The shield of Guiana is com were sent to specialists in the USA, England,
posed of rocks of the Precambrian Era, essentially France, the Netherlands, Belgium, Sweden, Co
of granites and gneisses. The soil corresponds to a lombia, and Brazil. Some of the samples, however,
red latosol. The annual precipitation is estimated are still in the process of being identified, although
at about 2000 mm or more, with a dry season of the inventory was finished approximately in 1967.
varying length between January and April. Special Most difficulties in the identification arose among
floristic studies were carried out around the camps the Burseraceae (the genus Protium), the Sapota
"Rio Grande", "EI Paraiso", and "EI Dorado". ceae (Pouteria), Myrtaceae, and Rosaceae (Lican
In the interior of the forest, some special edap ia). In spite of all the efforts of taxonomists, we
hic formations are included such as periodically still expect some misinterpretations and unidentif-
2
ied species. For these reasons, the vernacular cm DBH in the adult stage, such as some Rubia
names are still of great value. In the book" Struc ceae, Capparidaceae, and Melastomaceae. These
tural patterns of tropical barks" I tried to explain trees were also included, however, in an attempt
these names and to trace their origin (Roth to cover the maximum number of genera and spe
1981 ). cies.
The principal objective of the dendrological stu The collection of fruit and seed samples was
dies was the collection of wood samples for tech carried out by the forestry engineer Dr. Bernard
nological purposes. These samples were processed Rollet, who directed the inventory program of the
in the Instituto Forestal in Merida, Venezuela, and FAO. In total, about 56 different plant families
in the Instituto Technol6gico Forestal of the U ni with 370 species are included in these studies. As
versity of Merida. For further information the far as it was possible each family with its species is
reader is referred to B. Rollet "Inventario forestal described in the special part concerning fruit and
en la Guayana Venezolana", MAC, Informe No. seed morphology, inner structure and dispersal.
3, parte 2, Caracas 1967. Trees and shrubs were The great advantage of this collection is the exact
classified into three categories: knowledge of the number of individuals of each
I. Vc ry tall trees of 30 m height or more (cate species, genus and family. In this way it is possible
gory A), to make exact calculations of the number of indi
2. Trees between 10 m and 29 m in height (ca viduals and their dispersal types, of the distribu
tegory a), and tion of certain dispersal types in the different for
3. Small trees and shrubs less than 10m high est strata, of the number of individuals in homo
(category aa). geneous groups, and of the number of species per
Most samples were taken from trees with an aver hectare, to mention only a few of such possibili
age diameter between 10 em and I m at breast ties.
height. However, some trees do not even reach 20