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Dipterocarp Biology, Ecology, and Conservation
Dipterocarp
Biology, Ecology,
and Conservation

Jaboury Ghazoul
ETH Zurich

1
3
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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© Jaboury Ghazoul 2016
The moral rights of the author have been asserted
First Edition published in 2016
Impression: 1
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a retrieval system, or transmitted, in any form or by any means, without the
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contained in any third party website referenced in this work.
For Gnome, for ever.
 Acknowledgements
My family have patiently (and sometimes impa-
tiently) borne the brunt of my commitment to this
enterprise. I promise that I will not think (much)
about another book project for at least a few weeks.
My wife, Gnome Liston, deserves as much grati-
tude as I can ever muster, and more, not only in
helping me pull together and organize material for
this book, but, more importantly, in accommodat-
ing my self-imposed isolation to complete this book
in its final stages. In a bid to avoid any further fam-
ily strife, I also apologize to each of my children in
equal measure for not spending as much time with
them as I, and presumably they, would have liked in
the past few months. My son Alligin deserves extra
thanks for modelling dammar with such elegance.
A number of people have helped me with this
book. After my wife, two people stand out. Nora
Gallarotti spent many hours securing the necessary
permissions for the figures and illustrations repro-
duced in this book. A great deal of thanks are due to
her for getting on with the job when I was otherwise
preoccupied with the text. Alice Hughes has been
very generous with her time in drafting some of the
maps used in the book. She offered her help with
such willingness that it has been all too easy for me
to ask her to do just one more thing. She will receive
the promised beers in return, though this is scant
recompense for her efforts.
Many people have offered and provided photos,
and although I have not been able to use them all, I
am very grateful for these generous offers of assis-
tance. The photos I have used have come from Julia
Born, Rhett Butler, Min Cao, Richard Davies, Varun
Chopra, Arthur Chung, Aline Finger, Terry Henkel,
Sascha Ismail, Jin Jianhua, Marina Khaytarova, Bill
Laurance, Ch’ien Lee, Patrice Levang, James Mar-
grove, Cerlin Ng, Susan Page, Tony Rodd, Marco
Schmidt, Lee Su See, Uma Shankar, James Smith,
Thuy Nguyen Thi, and the Centre for International
Forestry Research. David Burslem, Richard Corlett,
Nimal Gunatilleke, Nourou Yorou, Chris Kettle,
and Chris Philipson provided additional informa-
tion, unpublished data, or comments, or indeed
helped shape the structure and content of the book.
Finally, this book would never have been written
were it not for the long standing collaborations I
have had with colleagues in Sabah, and particularly
at the Sepilok Forest Research Centre. Reuben Ni-
lus, Eyen Khoo, Chey Vun Khen, Robert Ong, and
Colin Maycock have long contributed to and sup-
ported our research on dipterocarps, without which
I would know nothing.
vii
Contents

1 The Dipterocarps  1
Introducing the Dipterocarpaceae  1
Economic and conservation contexts  3
Dipterocarp science  4
Book structure  6

2 Description and Identity  9

A short history of discovery and description  9
Taxonomy of the Dipterocarpaceae  10
Monotoideae  12
Pakaraimoideae  15
Dipterocarpoideae  15
Morphology and field identification  16
Tree form  18
Bark form and structure  19
Leaves  23
Flowers  26
Fruit  27
Forester classification  29
Conclusions  29

3 Distribution and Formations  33

Global distribution  33
The Asian dipterocarps  35
Dipterocarps in Africa  36
South American dipterocarps  39
Ecoregional distribution of Asian dipterocarps  40
A note on climate  42
A note on soils  43
Aseasonal formations  44
Seasonal evergreen formations  53
Deciduous and semi-deciduous formations  57
Other formations  62
The distributional limits of Asian dipterocarps  66
Conclusions  67

ix
x   C o n t e n t s

4 Evolution and Biogeography  69

Affiliation of Dipterocarpaceae within angiosperms  69
Phylogeny of the Dipterocarpoideae subfamily  70
The Dipterocarpeae tribe  72
The Shoreae tribe  72
Reconstructing the dipterocarp phylogeny  73
Evolutionary history and biogeography  73
Evolutionary origins  73
Out of Africa  75
Fossil evidence  76
Southeast Asian Miocene spread and radiation  78
Quaternary climatic and sea level changes  79
The Kangar–Pattani Line  82
East of Sundaland  82
Endemism  83
Widespread species  86
Sister species and hybrids  87
Conclusions  88

5 Reproduction and Population Genetics  89

General flowering and mast fruiting  89
Drivers of general flowering  94
Evolutionary origins of general flowering  96
Reproductive phenologies in seasonal zones  101
Pollination ecology  101
Pollination in Malesian dipterocarp forests  101
Pollination in seasonal dipterocarp forests  106
Fruit development  108
Seed dispersal  111
Population genetics and gene flow  114
Genetic diversity  114
Breeding system and outcrossing rates  118
Apomixis, polyembryony, and polyploidy  119
Hybridization  121
Seed predation  122
Insect pre-dispersal seed predation  122
Vertebrate pre- and post-dispersal predation  124
Conclusions  125

6 Light Acquisition  127

Light  128
Architecture  128
Light utilization  131
Relative growth rates  131
Sunflecks  133
Growth rate/survival trade-offs  135
Canopy gaps  137
Conclusions  146
 C o n t e n t s    xi

7 Water Relations  147

Drought  148
Drought avoidance and drought tolerance  148
Seed germination and seedling establishment  151
Seedlings and saplings  151
Trees  153
Continental comparisons  156
Too much water  158
Species distributions and water availability  160
Regional climate 160
Local topography  161
Conclusions  162

8 Nutrients  163
Soil types  163
Soil resources and dipterocarp distribution  164
Soil and forest structure  165
Broad-scale edaphic associations and diversity relations  165
Responses to nutrients  166
Spatial heterogeneity in nutrients  167
Interactions with light, topography, and herbivory  169
Resource partitioning and species coexistence  171
Conclusions  172

9 Population and Disturbance Dynamics  173

Tree mortality and gap formation  173
Dynamics of seedling populations  174
Competition among seedlings  174
Density and distance dependence  175
Adult architecture and phenology  178
Canopy stature and tree structure  178
Leaf phenology 179
Disturbance and dynamics  181
Drought  181
Fire  183
Storms and windthrow  184
Landslips  185
Episodic disturbances and forest dynamics  186
Conclusions  187

10 Community Ecology  189

Dipterocarp dominance  189
Community patterns of dipterocarp distribution  190
Regional scale  190
Local scale  191
Forest structure  195
Canopy and tree heights  195
xii   C o n t e n t s

Biomass and stem density  197

The role of soil  198
Biotic interactions: herbivory  200
Herbivory and plant defence  200
Niche segregation through herbivory  200
Biotic interactions: mycorrhiza  204
Mycorrhizal diversity  204
Function  206
Responses to disturbance  207
Regional implications of general flowering  208
Conclusions  209

11 Exploitation, Conservation, and Restoration  211

Regional land use change in Southeast Asia  211
Values of forest  213
Impacts and threats  214
Logging  214
Forest conversion to plantations and agriculture  219
Fire  219
Invasives  227
Logging and fragmentation: effects on recruitment  228
Genetic vulnerability  229
Conserving genetic diversity in managed forests  230
Non-timber forest products  231
Oleoresins  231
Dammar  232
Illipe nuts and butter fat  234
Camphor  234
Climate change  234
Conservation  235
Conservation status and protection  235
Conservation value of logged and secondary forests  238
Forest restoration and rehabilitation  240
Restoration  241
Ex situ propagation and conservation 245
Conclusions  247

Appendix 1 Timber Classifications of Various Dipterocarp Genera  249

Appendix 2 Key to the Main Dipterocarp Lowland Forest Types of Asia West of the Wallace Line  255
Appendix 3 Flowering Characteristics and Pollinators of Dipterocarp Species  257
References  261
Index  299
 A B

C D

Plate 1 Flowers of Hopea ponga (A), Vateria indica (B), Shorea bracteolata (C), and Dipterocarpus obtusifolius (D), showing the typical
overlapping (imbricate) corolla structure. Photos of H. ponga and V. indica by Sascha Ismail; photos of S. bracteolata and D. obtusifolius by Marina
Khaytarova. (See Figure 2.16 on page 27)
Plate 2 Dipterocarp species richness for terrestrial ecoregions of the Indo-Pacific. From Terrestrial Ecoregions of the Indo-Pacific by E. Wikamanayake, E. Dinerstein, C.J. Loucks et al. Copyright ©
2002 Island Press. Reproduced by permission of Island Press, Washington, DC. (See Figure 3.3 on page 37)
Plate 3 Dipterocarpaceae endemism for terrestrial ecoregions of the Indo-Pacific. From Terrestrial Ecoregions of the Indo-Pacific by E. Wikamanayake, E. Dinerstein, C.J. Loucks et al. Copyright ©
2002 Island Press. Reproduced by permission of Island Press, Washington, DC. (See Figure 4.7 on page 85)
 A B

Plate 4 Developing fruit of Parashorea malaanonan (A), and Hopea nervosa (B), at Danum Valley, Sabah, showing the photosynthesizing wings
prior to full maturation. Photos of P. malaanonan and H. nervosa by Ch’ien Lee, and D. lanceolata by James Smith. (See Figure 5.11 on page 109)

1.0 Species
S. acuminatissima
0.9 S. argentifolia
S. beccariana
0.8
S. falciferoides
S. gibbosa
0.7
H. beccariana
Proportion of fruit

0.6 D. humeratus
D. kerrii
0.5 S. macroptera
S. mecistopteryx
0.4 S. seminis
S. smithiana
0.3
S. xanthophylla

0.2

0.1

0.0

0 5 10 15 20 25 30 35
Distance dispersed (m)

Plate 5 Simulated dispersal kernals of 13 species based on release from height of 30 m and mean wind speed of 1.72 m s–1. From Smith et al.
(2015). (See Figure 5.16 on page 112)
Plate 6 An emergent Shorea johorensis at Danum Valley,
Plate 7 Olene mendosa (Lymantriidae) on a Shorea waltonii leaf (A),
Sabah, laden with mature fruit that can, potentially, be dispersed
and Setora sp. (Limacodidae) on a Shorea kudatensis leaf. Photos by
over several hundred metres, but more often fall within a few
Arthur Chung. (See Figure 10.3 on page 201)
tens of metres of the tree. Photo by Jaboury Ghazoul.
(See Figure 5.18 on page 114)

A B

Plate 8 Exopholis hypoleuca (Scarabaeoidea: Melolonthidae) on a Dryobalanops lanceolata leaf (A), and Hypomeces squamosus (Curculionidae)
on a dipterocarp leaf (B). Photos by Arthur Chung. (See Figure 10.4, on page 201)
 A B

Plate 9 Cypa decolor (Sphingidae) feeding on the flowers of Dryobalanops lanceolata (A), and Rapala manae ingana (Lycaenidae) on Shorea
argentifolia flowers (B). Photos by Arthur Chung. (See Figure 10.5 on page 201)

A 1973 FOREST COVER B FOREST COVER CLEARANCE (1973–2010)

1973 forest Forest clearance (1973–2010)

1973 non-forest

0 125 250 500

Km

C LOGGING ROADS (1973–2010) D 2010 INTACT, LOGGED FOREST

and PLANTATIONS
Primary logging roads (1973–2010)
Public roads 2010 logged forest
2010 intact forest
2010 industrial plantations

Plate 10 Four decades of forest persistence, clearance, and logging on Borneo. (A) Forest (dark green) and non-forest (white) in year 1973, and
residual clouds (cyan). (B) Areas of forest loss during 1973–2010 (red). (C) Primary logging roads from 1973 to 2010 (yellow lines). (D) Remaining
intact forest (dark green), remaining logged forest (light green), and industrial oil palm and timber plantations (black) in year 2010. From Gaveau
et al. (2014). (See Figure 11.1 on page 212)
 Main Areas and Causes of
Forest Change in SE Asia
Agriculture
Coffee/Tea/Rubber/Other Crops
Oil Palm
Oil Palm+Tree Plantations
Logging
Logging /Shifting Cult./Agriculture
Tree Plantations/Agriculture
Rubber/Tree Plantations /Logging
Rubber
Small Scale Change
Afforestation, Reforestation, Regrowth

0 250 500 Kilometers

Plate 11 Regional pattern of main areas and causes of forest change in Southeast Asia. From Stibig et al. (2014). (See Figure 11.2 on page 213)
 Anisoptera (Mersawa) Cotylelobium (Resak) Dipterocarpus (Keruing) Dryobalanops (Kapur)

Hopea (Merawan) Hopea (Giam) Parashorea (Gerutu) Shorea section Anthoshorea

(White Meranti)

Shorea section Mutica Shorea section Mutica Shorea section Richetia Shorea section Pachycarpae
(Dark Red Meranti) (Light Red Meranti) (Yellow Meranti) (Melantai)

Shorea section Rubroshorea Shorea section Shorea

Shorea albida (Alan Batu) Shorea albida (Alan Bunga)
(Red Balau) (Balau)

Shorea uliginosa Neobalanocarpus

(Meranti Bakau) (Chengal)

Plate 12 (See Figure Appendix 1 on page 253)

 C H A PT ER 1
The Dipterocarps
To the delight of foresters, Southeast Asia’s low-
land rain forests are largely dominated by trees of
a single family, the Dipterocarpaceae, long valued
for their timber qualities and exploited accordingly.
Dipterocarp-dominated forests stretch from India
across continental Southeast Asia into the Malesian
archipelago where they reach their greatest species
number. Here, biodiversity as a whole achieves a
richness and variety equivalent to that of the Ama-
zon and unsurpassed elsewhere. The competing
demands of human exploitation of forest resources
and those of biodiversity conservation have long
been played out among Asia’s forested regions. The
Dipterocarpaceae, as the dominant plant group but
also the principal resource, has for decades been at
the centre of this conflict.
By the turn of the twenty-first century the battle to
preserve the lowland forests of Southeast Asia had
largely been lost to exploitation and land clearance.
Yet forest patches of varying sizes remain, protected
areas have been established, and restoration of de-
graded forests is underway. The future success of
this rearguard action will depend on good use of
the accumulated scientific knowledge on diptero-
carps, but also on society’s recognition of the so-
cial and ecological values provided by dipterocarp
forests and the biodiversity they contain. There are
positive signs that society is indeed increasingly ac-
knowledging the value of Asian tropical biodiver-
sity, even as it is being lost, as well as the variety of
ecosystem services that Asian forests provide. This
bodes well for the future recovery and conservation
of dipterocarp forests, even if in the short to medium
term many political and scientific challenges remain.
I can contribute but little to the political dialogue
about land use, resource exploitation, and forest
Dipterocarp Biology, Ecology, and Conservation. Jaboury Ghazoul. © Jaboury Ghazoul 2016.
Published 2016 by Oxford University Press. DOI 10.1093/acprof:oso/9780199639656.001.0001
conservation. Rather, my intention with this book
is to collate scientific knowledge on the diptero-
carp family and dipterocarp forests to provide the
foundation for future management and restoration
work. As an ecologist I also aim to synthesize the
state of ecological knowledge on this important
plant family. The sheer diversity and abundance of
dipterocarps in tropical Asia’s forests ensure that
any study of forest ecology and biodiversity in this
region would be incomplete without understand-
ing the Dipterocarpaceae. Hence I hope that this
book will additionally provide the necessary foun-
dation for continued understanding of this fascin-
ating biome.
Introducing the Dipterocarpaceae
The Dipterocarpaceae are subcanopy, canopy, or
emergent trees, with many species exceeding 50 m
in height (Figure 1.1). Asian dipterocarp rain forests
have the highest stature of any tropical forest. The
tallest known flowering tree in the tropics, meas-
ured at 88.3 m, is a dipterocarp, Shorea faguetiana in
Tawau Hills National Park, Sabah, Malaysia. This
is almost as high as the tallest tree ever recorded in
the tropics, the gymnosperm Araucaria hunsteinii in
Papua New Guinea, which in 1941 was measured at
88.9 m by a climber carrying a steel tape to the top
(L. G. Cavanaugh quoted by Gray 1975). Records
from the early twentieth century indicate trees of
massive size. The girths of two trees on the Malay
Peninsula, Neobalanocarpus heimii and Dryobalanops
sumatrana, were measured at 12 m, almost 4 m diam-
eter (Foxworthy 1927). Such trees no longer exist.
It is not so much size that has made dipterocarps
attractive to foresters, but rather their abundance.
2   Th e D ip t e r o carp s
Figure 1.1 A tall canopy emergent Parashorea malaanonan tree at
Danum Valley, Sabah. Photo by Jaboury Ghazoul.
Ecologists frequently speak of Asian lowland forests
as being ‘dominated’ by the Dipterocarpaceae (see
Chapter 10 for a discussion of the possible causes
of family dominance), as I have in the opening
sentence of this book, but what does this actually
mean? In broad terms, the family Dipterocarpaceae
comprises between 5 and 20% of all tree species at
any particular location within most Asian lowland
forests (Chapter 3). In Borneo, where the diptero-
carps attain their highest diversity, they account for
21.9% of all trees based on data from 28 lowland
forest sites (Slik et al. 2003). The genus Shorea alone
comprises 12.3% of trees. While this in itself dem-
onstrates that dipterocarps are disproportionally
represented among tree families in Asian forests,
a further striking feature is that they are, predomi-
nantly, large canopy trees. Thus dipterocarps ac-
count for 20–50% (or more) of forest basal area, and
often well over 50% of canopy trees (Ashton 1982,
1988b) (Table 1.1; see also Chapter 3). No other trop-
ical biome is so well represented, or has its ecology
shaped to such a degree, by a single plant family
(although the legume family, and particularly the
Caesalpinioideae, is similarly dominant in tropical
Africa and America). Indeed, the abundance of dip-
terocarps in Asian tropical forests has far reaching
implications for the community composition and
forest structure, and for the history of silviculture
and subsequent land development of the region.
The Dipterocarpaceae includes around 500 tree
species with a pantropical distribution, although
the large majority are located in Southeast Asia.
The early development of several Asian nations
was built upon dipterocarp timber that has long
been the mainstay of the global tropical timber
trade. In recent decades unsustainable exploitation
of timber, and forest clearance for agriculture, has
caused massive losses of dipterocarp forests, and
many dipterocarp species are now listed as en-
dangered, and all too often critically so. While this
in itself is of concern, there are far wider implica-
tions for loss of global biodiversity, the richness of
which within dipterocarp forests is second only
to the forests of South America. Moreover, many
features of dipterocarp forests mark them out as
being ecologically unique. These forests are home
to orang-utan, the only non-African ape, and an
unusual variety of volant vertebrates, to name but
two examples (for a full description of the gen-
eral biodiversity and character of Asian forests see
Ghazoul and Sheil 2010; Primack and Corlett 2005).
The cycles of resource abundance and scarcity,
driven by dipterocarp reproductive biology in the
aseasonal tropics, are also unprecedented in other
tropical regions.
Table 1.1 Percentage of dipterocarps among very large canopy and
emergent trees (exceeding 75 cm DBH) in the aseasonal lowland
dipterocarp rain forests.
Peninsular Malaysia <57
Sarawak 76
Sabah 78–98
Philippines 100
After Whitmore (1984b).

Figure 1.2 An oil palm plantation on land that was formerly lowland dipterocarp forest in Sabah, Borneo. Photo by Jaboury Ghazoul.
Economic and conservation contexts
It is the value of dipterocarp timber that was the
impetus for the early colonial exploration of dip-
terocarp forests, and their resulting exploitation
(Boomgaard et al. 1996). The Dutch East India
Company first exploited timber in Java upon its
annexation of Jakarta in 1619. The primary use of
timber at that time was shipbuilding. Formal at-
tempts to manage forest resources in the region did
not emerge until the early 1800s on the colonization
of Indonesia by the Dutch republic. A Forest Board
was established in 1808, and this regulated har-
vesting, forest tenure, and silvicultural practices.
After the colonial period, the extraction of timber
from these forests became the foundation for the
economic development of the independent nation
states of Indonesia and Malaysia. In the 1970s and
1980s timber from natural forests accounted for
more than 50% of income of the Malaysian states
Ec o n o mic a n d c o n s e rvat i o n c o n t e x t s    3
of Sabah and Sarawak. Rates of extraction outpaced
rates of regrowth of commercial species, and second
or third harvests have become successively less eco-
nomically rewarding. This has reduced the oppor-
tunity costs of forest conversion to other land uses.
Indeed, regional economic growth and population
expansion have since driven agricultural expansion
and the conversion of logged and degraded forests
to, for example, rubber, acacia, and oil palm plant-
ations (Figure 1.2). Other forms of resource exploit-
ation within dipterocarp forests include mining,
while construction of roads and dams provides the
access and energy needed for the opening up and
development of many remote forest regions. Much
of this development undoubtedly materially bene-
fits people living in these regions, even if it has ac-
celerated the loss of dipterocarp forests.
Now, in 2016, few tracts of lowland dipterocarp
forests remain outside protected areas, and those
that do are either degraded or fragmented, or both.
4   Th e D ip t e r o carp s
In Peninsular Malaysia for example, the only re-
maining extent of relatively undisturbed diptero-
carp forest lies within Taman Negara, a 4343-km²
protected area that is surrounded on all sides by
land devoid of any natural forest. Elsewhere in Ma-
laysia, remaining lowland forest patches are one,
or even two, orders of magnitude smaller. The 438-
km2 Danum Valley Conservation Area and the 43-
km2 Sepilok Forest Reserve are similarly bordered
by degraded forest or agricultural lands. In recog-
nition of their value for biodiversity, and their eco-
nomic benefits through ecotourism, many of these
remaining forest patches are now afforded some
protection by various conservation designations.
Increasing scarcity of unlogged dipterocarp for-
est has weakened the value of the forest sector. For-
estry (harvesting and silviculture) still contributes
US$5.1 billion to Indonesia’s economy, but this rep-
resents just 1% of current GDP (ITS Global 2011).
Timber extraction has been overtaken by wood and
wood manufacturing and pulp and paper produc-
tion (which together contribute 2.6% to Indonesia’s
GDP), largely through the establishment of exotic
tree plantations on former dipterocarp forests.
The loss of lowland dipterocarp forests does far
more than undermine the viability and future of the
timber industry. Indeed the global value of these for-
ests is in the immense and unique biodiversity they
contain. National benefits include revenues from
one of the fastest growing industries w ­ orldwide—
that of tourism which, in Southeast Asia, is to a large
extent built upon the beauty and biodiversity of the
natural environment. In Malaysia in 2011 tourism
employed 14% of the workforce and generated
revenues worth MR36.9 billion (US$12.3 billion at
2011 exchange rates) to GDP from foreign visitors,
a large share of which is attracted by Malaysia’s
forest parks and wildlife (PEMANDU 2010). By
comparison, timber and wood products accounted
for MR20.5 billion in 2010 (US$6.8 billion), and this
sector does not even feature in the 12 National Key
Economic Areas identified by the Malaysian gov-
ernment in 2010.
Increasing confidence in an incipient market for
carbon, buoyed by commitments from some coun-
tries to finance carbon conservation through pro-
tection of natural forests (e.g. Norway’s pledge
of US$1 billion to Indonesia for forest protection
through REDD+), has also focused attention on
Asia’s remaining dipterocarp forests, which have
some of the highest above-ground biomass values
of any habitat.
The combination of environmental tourism and
carbon conservation, coupled with continuing pres-
sure to deliver sustainable timber production, has
motivated governments to designate protected ar-
eas and incentivized forest restoration efforts across
tropical Asia. The security of protection afforded
to forest reserves is, however, far from certain.
Governments must balance multiple priorities, in-
cluding development of sectors that might be det-
rimental to forest cover. While these are largely
issues of policy and economics, their implications
encompass conservation, carbon emissions, and
land management.
Forest restoration efforts, meanwhile, have had
to grapple with several challenges in the context
of Asia’s dipterocarp forests. These challenges are
largely ecological in nature and relate to, for exam-
ple, securing and maintaining viable seed, match-
ing species to suitable habitat conditions, and
maintaining genetic diversity. The security of Asia’s
dipterocarp forests, and the biodiversity they con-
tain, is also exposed to future climate change. Fu-
ture climate trajectories for Southeast Asia remain
uncertain, but some fear a trend of increasing fre-
quency and intensity of El Niño events, which drive
droughts and facilitate forest fires.
Dipterocarp science
Research interest in dipterocarp forests has it-
self evolved. It was the silviculturalists who first
showed interest in dipterocarp forests on account
of their timber value, and early studies empha-
sized such interests. Ecologists followed the for-
esters, and sought to understand dipterocarp
biology and the wider functioning of dipterocarp
forests. New research interest in the past decade
or so has been spurred by the potential of diptero-
carp forests to store carbon, either as biomass or
as soil carbon, and the need to better manage for-
ests. ­Conservation-minded researchers, who have
sought to understand the implications of rapid for-
est loss on biodiversity, see carbon values as an op-
portunity to preserve remaining forest stands and

restore degraded stands. Now we are developing
knowledge for forest restoration activities, and are,
to a large extent, entering new scientific territory.
Yet restoration is perhaps the best long-term hope
for retaining much of Asia’s biodiversity while miti-
gating greenhouse gas emissions.
The history of scientific discovery reflects the
commercial interest of dipterocarps to early colonial
foresters and associated botanists. Early work on
dipterocarps naturally focused on the Malay Archi-
pelago where European colonial centres were estab-
lished and where dipterocarps are most diverse. Yet
it is easy to be distracted by the impressive stature
and diversity of insular Southeast Asia’s lowland
rain forests and its dipterocarps. We should rec-
ognize that the family is distributed across a wide
range of other climatic and biogeographic realms
in continental Southeast Asia. Also easily forgotten
are representatives of the family, fewer in number
and of smaller stature, in Africa and South America.
Such patterns of abundance and distribution raise
interesting questions for evolutionary biologists,
biogeographers, and ecologists alike.
It is possible that after many decades of research,
the Dipterocarpaceae might be the best known trop-
ical plant family. New tools from genetics, remote
sensing, and modelling, more refined experimental
and analytical work, and valuable data from long-
term sample plots continue to expand and shape
our understanding of dipterocarps, dipterocarp
forests, and tropical forests more generally. Yet des-
pite being the most intensively researched tropical
tree family, knowledge on dipterocarps mostly lies
within often obscure taxonomic treatise, technical
academic theses and journal papers, or inaccessible
forestry reports. Substantial efforts have been made
to consolidate information on taxonomy (Ashton
1982; Newman et al. 1995, 1996a, b, 1998a, b, c, d;
Symington 2004) and silviculture (Appanah and
Turnbull 1998; Wyatt-Smith 1965), but progress on
the ecology and evolution of the family has out-
paced such treatment.
Science, and particularly ecological research, has
an important contribution to make in improving fu-
ture forest policy, management, and restoration. In
pursuing such goals, it would seem foolish for trop-
ical forest ecologists to focus research attention on a
single plant family. Yet in the case of tropical Asian
D ip t e r o carp s ci e n c e    5
forests such an approach might be at least partially
justified. After all, many tropical Asian forests, and
certainly those most valuable from economic and
conservation perspectives, are characterized by
the great abundance and diversity of the Diptero-
carpaceae among their large canopy-forming trees.
The wildlife dynamics of tropical Asian forests is
also heavily influenced by particular features of
dipterocarp reproductive ecology (Chapter 5). Dip-
terocarp biomass is disproportionate to that of other
families, and while dipterocarp trees have been the
main target for timber extraction, they have also
been used extensively for restoration. Any ecologist
seeking to understand the dynamics of Asian trop-
ical rainforests would, of course, be remiss to focus
only on the Dipterocarpaceae, but an understand-
ing of this single family certainly goes a long way
towards a comprehensive understanding.
The purpose of this book is to synthesize know-
ledge on the ecology of the Dipterocarpaceae, taking
account of the considerable recent work in multiple
branches of ecological science. I have intentionally
restricted the scope to the Dipterocarpaceae, and I
avoid the temptation to regale readers with tales
(scientific or otherwise) of the wider richness and
ecology of Asian tropical forests, except when it is
directly relevant to dipterocarp ecology. Other re-
cent texts fulfil this role (Ashton 2014; Corlett 2014;
Ghazoul and Sheil 2010; Primack and Corlett 2005).
I hope that this synthesis of the Dipterocarpaceae
will provide an up-to-date scientific foundation for
the management of the remaining dipterocarp for-
ests, and a basis for future dipterocarp restoration
efforts. I caution, however, that as an ecologist I
write as an ecologist, and my treatment of subjects
will necessarily reflect my professional perspective.
I make little effort to detail silvicultural practices or
management approaches beyond their relevance
to ecology and conservation. There may be a need
for a comprehensive treatment of dipterocarp for-
est management systems, but I am not the one to
write it.
Moreover, as an ecologist I am fascinated by
some of the more theoretical ecological questions to
which research on dipterocarps has made substan-
tial contributions. This will be evident in my treat-
ment of the literature. As an academic (or armchair)
conservationist, I also hope that this book will
6   Th e D ip t e r o carp s
motivate and inform continued efforts to conserve
dipterocarps forests and the remarkable biodiver-
sity they contain.
I justify the effort expended on this book through
its purpose to provide forest managers and re-
searchers with a synthesis of available information
on Dipterocarpaceae that is necessary to prepare
them for future research and management chal-
lenges. These challenges are likely to be very dif-
ferent to those of the past century, and any aspiring
scientist or manager needs knowledge and skills
that go far beyond what I can present in this book.
Nonetheless, the biological and ecological central-
ity of dipterocarps in Asian tropical forests and the
centrality of tropical forests for global issues of bio-
diversity, carbon sequestration, and resource use
are reasons enough for a volume on this important
plant family.
A final caveat: While I have already acknow-
ledged that dipterocarps are a pantropical family,
this book unavoidably focuses on the Asian topics
where the vast majority of dipterocarps occur. In-
deed, the ecology of tropical Asian forests is inex-
tricable from the ecology of the Dipterocarpaceae,
whereas in other tropical regions dipterocarps are
merely peripheral players.
India
Indo-Burma
Sri Lanka
Sundaland
Figure 1.3 Bioregions of South and Southeast Asia, as referred to in this book. Map by Alice Hughes.
Book structure
The book is divided into two sections. The first sec-
tion begins by describing the Dipterocarpaceae and
the characteristics of its component genera (Chap-
ter 2). The distribution and diversity of dipterocarp
forest formations is detailed in Chapter 3. In this
chapter, and indeed throughout the book, I refer to
named biogeographic units. There is not complete
agreement on the definition of these names so, for
clarity, the biogeographical terms I use in this book
are illustrated in Figure 1.3. The need for a well-
resolved phylogeny is key for a fuller understand-
ing of dipterocarp ecology and evolution. Although
this remains an incomplete task, it is addressed in
Chapter 4.
The second part of the book focuses on the ecol-
ogy of the Dipterocarpaceae. The reproductive biol-
ogy of the family has long been a subject of interest,
and is elaborated in Chapter 5. Three chapters on
ecophysiology follow, tackling different strategies
for the acquisition of light, water, and nutrients
(Chapters 6–8). The common theme of these chap-
ters is the different life history strategies that serve
to underpin niche segregation among dipterocarps.
Most of this work has been undertaken in the wet
Philippines
New Guinea
Wallacea

aseasonal tropics, and it is the immense species di-
versity of forests in this region, and our attempts
to understand the mechanisms for species coexist-
ence, that provide the fundamental inspiration that
lies behind much of this research. Apart from con-
tributing to fundamental ecological questions re-
lating to species coexistence, these topics also hold
much interest for management of forest stands and
ecological restoration. The accumulated ecophysio-
logical knowledge provides a basis for understand-
ing dipterocarp population biology and community
structure (Chapters 9 and 10), particularly in rela-
tion to interactions with other organisms, be they
mycorrhizas, herbivores, or seed predators.
B o o k s t r u c t u r e    7
The book concludes with a final chapter on con-
servation and management of dipterocarps and
dipterocarp forests. Whole books can be written
on this topic, and indeed have been (Corlett 2014;
Ghazoul and Sheil 2010; Schulte and Schöne 1996;
Sodhi and Brook 2006; Whitmore 1984b). It is not
my intention to recapitulate what has been said be-
fore. Rather, in Chapter 11, I provide a short over-
view of conservation and management issues as
they particularly pertain to dipterocarps and dip-
terocarp forests. The book concludes with a syn-
thesis of prospects for the future, and how ecology
might inform conservation and good management
of dipterocarp forests.
Another random document with
no related content on Scribd:
otra alguna,
ueys vna perfeçion jamas
oyda,
ueys una discreçion, qual fue
ninguna,
de hermosura y graçia
guarnescida?
¿ueys la que está domando a
la fortuna
y a su pesar la tiene alli
rendida?
la gran doña Leonor Manuel
se llama,
de Lusitania luz que al orbe
inflama.
Doña Luisa Carrillo, que en
España
la sangre de Mendoça ha
esclareçido:
de cuya hermosura y graçia
extraña,
el mismo amor, de amor está
uençido,
es la que a nuestra Dea ansi
acompaña
que de la uista nunca la ha
perdido:
de honestas y hermosas claro
exemplo,
espejo y clara luz de nuestro
templo.
¿Ueys una perfeçion tan
acabada
de quien la misma fama está
embidiosa?
¿ueys una hermosura más
fundada
en graçia y discreçion que en
otra cosa,
que con razon obliga a ser
amada
porque es lo menos de ella el
ser hermosa?
es doña Eufrasia de Guzman
su nombre,
digna de inmortal fama y gran
renombre.
Aquella hermosura
peregrina
no uista en otra alguna sino en
ella,
que a qualquier seso apremia
y desatina,
y no hay poder de amor que
apremie el della,
de carmesí uestida y muy más
fina
de su rostro el color que no el
de aquella,
doña Maria de Aragon se
llama,
en quien se ocupará de oy
más la fama.
¿Sabeys quién es aquella
que señala
Diana, y nos la muestra con la
mano,
que en graçia y discreçion a
ella yguala,
y sobrepuja a todo ingenio
humano,
y aun ygualarla en arte, en ser
y en gala,
sería (segun es) trabajo en
uano?
doña Ysabel Manrique y de
Padilla,
que al fiero Marte uenze y
marauilla.
Doña Maria Manuel y doña
Ioana
Osorio, son las dos que estays
mirando
cuya hermosura y graçia sobre
humana,
al mismo Amor de amor está
matando:
y esta nuestra gran Dea muy
vfana,
de ueer a tales dos de nuestro
uando,
loallas, segun son es
escusado:
la fama y la razon ternan
cuydado.
Aquellas dos hermanas tan
nombradas
cada una es una sola y sin
segundo,
su hermosura y graçias
extremadas,
son oy en dia un sol que
alumbra el mundo,
al biuo me paresçen
trasladadas,
de la que a buscar fuy hasta el
profundo:
doña Beatriz Sarmiento y
Castro es una
con la hermosa hermana qual
ninguna.
 El claro sol que ueys
resplandeçiendo
y acá, y allá sus rayos ya
mostrando,
la que del mal de amor se está
riendo,
del arco, aljaua y flechas no
curando,
cuyo diurno rostro está
diziendo,
muy más que yo sabré dezir
loando,
doña Ioana es de Çarate, en
quien vemos
de hermosura y graçia los
extremos.
Doña Anna Osorio y Castro
está cabe ella
de gran valor y graçia
acompañada,
ni dexa entre las bellas de ser
bella,
ni en toda perfeçion muy
señalada,
mas su infelize hado vsó con
ella
de una crueldad no vista ni
pensada,
porque al ualor, linaje y
hermosura
no fuesse ygual la suerte, y la
uentura.
Aquella hermosura
guarnecida
de honestidad, y graçia sobre
humana,
que con razon y causa fue
escogida
por honra y prez del templo de
Diana,
contino uençedora, y no
uençida
su nombre (o Nimphas) es
doña Iuliana,
de aquel gran Duque nieta y
Condestable,
de quien yo callaré, la fama
hable[1256].
Mirad de la otra parte la
hermosura
de las illustres damas de
Valençia,
a quien mi pluma ya de oy
mas procura
perpetuar su fama y su
excelençia:
aqui, fuente Helicona, el agua
pura
otorga, y tú, Minerua, enpresta
sçiençia,
para saber dezir quién son
aquellas
que no hay cosa que ver
despues de vellas.
Las cuatro estrellas ved
resplandesçientes
de quien la fama tal ualor
pregona
de tres insignes reynos
desçendientes,
y de la antigua casa de
Cardona,
de la vna parte Duques
exçelentes,
de otra el trono, el sçeptro, y la
corona,
del de Segorbe hijas, cuya
fama
del Borea al Austro, al Euro se
derrama.
La luz del orbe con la flor de
España,
el fin de la beldad y
hermosura,
el coraçon real que le
acompaña,
el ser, valor, bondad sobre
natura,
aquel mirar que en verlo
desengaña,
de no poder llegar alli criatura:
doña Anna de Aragon se
nombra y llama,
a do por el amor, cansó la
fama.
Doña Beatrix su hermana
junto della
vereys, si tanta luz podeys
miralla:
quien no podré alabar, es sola
ella,
pues no ay podello hazer, sin
agrauialla:
a aquel pintor que tanto hizo
en ella,
le queda el cargo de poder
loalla,
que a do no llega
entendimiento humano
llegar mi flaco ingenio, es muy
en vano.
 Doña Françisca d'Aragon
quisiera
mostraros, pero siempre está
escondida:
su vista soberana es de
manera,
que a nadie que la vee dexa
con vida:
por esso no paresçe. ¡Oh
quién pudiera
mostraros esta luz, que al
mundo oluida,
porque el pintor que tanto hizo
en ella,
los passos le atajó de
meresçella.
A doña Madalena estays
mirando
hermana de las tres que os he
mostrado,
miralda bien, uereys que está
robando
a quien la mira, y biue
descuydado:
su grande hermosura
amenazando
está, y el fiero amor el arco
armado,
porque no pueda nadie, ni aun
miralla,
que no le rinda o mate sin
batalla.
Aquellos dos luzeros que a
porfia
acá, y allá sus rayos uan
mostrando,
y a la exçelente casa de
Gandia,
por tan insigne y alta
señalando,
su hermosura y suerte sube oy
dia
muy más que nadie sube
imaginando:
¿quién uee tal Margareta y
Madalena,
que tema del amor la horrible
pena?
Quereys, hermosas
Nimphas, uer la cosa,
que el seso más admira y
desatina?
mirá una Nimplia más que el
sol hermosa,
pues quién es ella, o él jamas
se atina:
el nombre desta fenix tán
famosa,
es en Valençia doña Cathalina
Milan, y en todo el mundo es
oy llamada
la más discreta, hermosa y
señalada.
Alçad los ojos, y vereis de
frente
del caudaloso rio y su ribera,
peynando sus cabellos, la
exçelente
doña Maria Pexon y
Çanoguera
cuya hermosura y gracia es
euidente,
y en discreçion la prima y la
primera:
mirad los ojos, rostro
cristallino,
y aquí puede hazer fin uuestro
camino.
Las dos mirad que están
sobrepujando,
a toda discreçion y
entendimiento,
y entre las más hermosas
señalando
se uan, por solo vn par, sin par
ni cuento,
los ojos que las miran
sojuzgando:
pues nadie las miró que biua
essento:
¡ued qué dira quien alabar
promete
las dos Beatrizes, Vique y
Fenollete!
Al tiempo que se puso alli
Diana,
con su diuino rostro y
excelente
salió un luzero, luego una
mañana
de Mayo muy serena y
refulgente:
sus ojos matan y su uista
sana,
despunta alli el amor su flecha
ardiente,
su hermosura hable, y
testifique
ser sola y sin ygual doña Anna
Vique.
Bolued, Nimphas, uereys
doña Teodora
Carroz, que del valor y
hermosura
la haze el tiempo reyna y gran
señora
de toda discreçion y graçia
pura:
qualquiera cosa suya os
enamora,
ninguna cosa nuestra os
assegura,
para tomar tan grande
atreuimiento,
como es poner en ella el
pensamiento.
Doña Angela de Borja
contemplando
uereys que está (pastores) en
Diana,
y en ella la gran dea está
mirando
la graçia y hermosura
soberana:
Cupido alli a sus pies está
llorando,
y la hermosa Nimpha muy
ufana,
en uer delante della estar
rendido
aquel tyrano fuerte y tan
temido.
De aquella illustre cepa
Çanoguera,
salio una flor tan extremada y
pura,
que siendo de su edad la
primauera,
ninguna se le yguala en
hermosura:
de su excelente madre es
heredera,
en todo quanto pudo dar
natura,
y assi doña Hieronyma ha
llegado
en graçia y disceçion al sumo
grado.
¿Quereys quedar (o
Nimphas) admiradas,
y uer lo que a ninguna dió
uentura:
quereys al puro extremo uer
llegados
ualor, saber, bondad y
hermosura?
mirad doña Veronica
Marradas,
pues solo uerla os dize y
assegura
que todo sobra, y nada falta
en ella,
sino es quien pueda (o piense)
meresçella.
Doña Luysa Penarroja
uemos
en hermosura y graçia más
que humana,
en toda cosa llega los
estremos,
y a toda hermosura uençe y
gana:
no quiere el crudo amor que la
miremos
y quien la uió, si no la uee, no
sana:
aunque despues de uista el
crudo fuego
en su vigor y fuerça buelue
luego.
Ya ueo, Nimphas, que
mirays aquella
en quien estoy continuo
contemplando,
los ojos se os yran por fuerça
a ella,
que aun los del mismo amor
está robando:
mirad la hermosura que ay en
ella,
mas ued que no çegueys
quiçá mirando
a doña Ioana de Cardona,
estrella
que el mismo amor está
rendido a ella.
Aquella hermosura no
pensada
que ueys, si uerla cabe en
nuestro uaso:
aquella cuya suerte fue
estremada
pues no teme fortuna, tiempo
o caso,
aquella discreçion tan
leuantada,
aquella que es mi musa y mi
parnaso:
Ioanna Anna, es Catalana, fin
y cabo
de lo que en todas por
estremo alabo.
 Cabe ella está un estremo
no uicioso,
mas en uirtud muy alto y
estremado,
disposiçion gentil, rostro
hermoso,
cabellos de oro, y cuello
delicado,
mirar que alegra, mouimiento
ayroso,
juyzio claro y nombre
señalado,
doña Angela Fernando, aquien
natura
conforme al nombre dio la
hermosura.
Vereys cabe ella doña
Mariana,
que de ygualalle nadie está
segura;
miralda junto a la exçelente
hermana,
uereys en poca edad gran
hermosura,
uereys con ella nuestra edad
ufana,
uereys en pocos años gran
cordura,
uereys que son las dos el
cabo y summa
de quanto dezir puede lengua
y pluma.
Las dos hermanas Borjas
escogidas,
Hippolita, Ysabel, que estays
mirando,
de graçia y perfeçion tan
guarnesçidas,
que al sol su resplandor está
çegando,
miraldas y uereys de quantas
uidas
su hermosura siempre ua
triumphando:
mirá los ojos, rostro, y los
cabellos,
que el oro queda atras y
passan ellos.
Mirad doña Maria
Çanoguera,
la qual de Catarroja es oy
señora,
cuya hermosura y graçia es de
manera,
que a toda cosa uençe y la
enamora:
su fama resplandeçe por do
quiera
y su uirtud la ensalça cada
hora,
pues no ay qué dessear
despues de uella,
¿quién la podrá loar sin
offendella?
Doña Ysabel de Borja está
defrente
y al fin y perfeçion de toda
cosa,
mira la graçia, el ser, y la
exçelente
color más biua que purpurea
rosa,
mirad que es de uirtud y graçia
fuente,
y nuestro siglo illustre en toda
cosa:
al cabo está de todas su
figura,
por cabo y fin de graçia y
hermosura.
La que esparzidos tiene sus
cabellos
con hilo de oro fino atras
tomados,
y aquel diuino rostro, que él y
ellos
a tantos coraçones trae
domados,
el cuello de marfil, los ojos
bellos,
honestos, baxos, uerdes, y
rasgados,
doña Ioana Milan por nombre
tiene,
en quien la uista pára y se
mantiene,
Aquella que alli ueys, en
quien natura
mostró su sçiençia ser
marauillosa,
pues no ay pasar de alli en
hermosura,
no ay más que dessear a una
hermosa:
cuyo ualor, saber, y gran
cordura
leuantarán su fama en toda
cosa,
doña Mençia se nombra
Fenollete,
 a quien se rinde amor y se
somete.

La cançion del çelebrado Orpheo,

fue tan agradable a los oydos de
Felismena, y de todos los que la
oyan, que assi los tenia
suspensos, como si por ninguno
de ellos uuiera passado más de lo
que presente tenian. Pues
auiendo muy particularmente
mirado el rico aposento, con
todas las cosas que en él auia
que uer, salieron las Nymphas por
una puerta de la gran sala, y por
otra de la sala a un hermoso
jardin, cuya uista no menos
admiraçion les causó que lo que
hasta alli auian uisto, entre cuyos
arboles y hermosas flores auia
muchos sepulchros de nimphas y
damas, las quales auian con gran
limpieça conseruado la castidad
deuida a la castissima diosa.
Estauan todos los sepulchros
coronados de enredosa yedra,
otros de olorosos arrayhanes,
otros de uerde laurel. De más
desto auia en el hermoso jardin
muchas fuentes de alabastro,
otras de marmol jaspeado, y de
metal, debaxo de parrales, que
por ençima de artifiçiosos arcos
estendian todas sus ramas, los
myrthos hazian cuatro paredes
almenadas, y por ençima de las
almenas, paresçian muchas flores
de jazmin, madreselua, y otras
muy apazibles a la uista. En
medio del jardin estaua una
piedra negra, sobre quatro pilares
de metal, y en medio de ella un
sepulchro de jaspe, que quatro
Nimphas de alabastro en las
manos sostenian, entorno dél
estauan muchos blandones, y
candeleros de fina plata, muy bien
labrados, y en ellos hachas
blancas ardiendo. En torno de la
capilla auia algunos bultos de
caualleros, otros de marmol
jaspeado, y de otras diferentes
materias. Mostrauan estas figuras
tan gran tristeza en el rostro, que
la pusieron en el coraçon de la
hermosa Felismena, y de todos
los que el sepulchro veyan. Pues
mirandolo muy particularmente,
vieron que a los pies dél, en una
tabla de metal que una muerte
tenía en las manos, estaua este
letrero:

Aqui reposa doña Catalina

de Aragon y Sarmiento cuya
fama,
al alto çielo llega, y se
auezina,
y desde el Borea al Austro se
derrama:
matéla, siendo muerte, tan
ayna,
por muchos que ella ha
muerto, siendo dama,
 acá está el cuerpo, el alma
allá en el çielo,
que no la meresçio gozar el
suelo.

Despues de leydo el Epigramma,

vieron cómo en lo alto del
sepulchro estaua vna aguda de
marmol negro, con vna tabla de
oro en las vñas, y en ella estos
uersos.

Qual quedaria (o muerte) el

alto çielo
sin el dorado Apollo y su
Diana
sin hombre, ni animal el baxo
suelo,
sin norte el marinero en mar
insana,
sin flor, ni yerua el campo y sin
consuelo,
sin el roçio d'aljofar la
mañana,
assi quedó el ualor, la
hermosura,
sin la que yaze en esta
sepultura.

Quando estos dos letreros

vuieron leydo, y Belisa entendido
por ellos quién era la hermosa
Nimpha que alli estaua sepultada,
y lo mucho que nuestra España
auia perdido en perdella,
acordandosele de la temprana
muerte del su Arsileo, no pudo
dexar de dezir con muchas
lagrimas: Ay muerte, quán fuera
estoy de pensar, que me as de
consolar con males agenos!
Dueleme en estremo lo poco que
se gozó tan gran ualor y
hermosura como esta Nimpha me
dizien que tenía, porque ni estaua
presa de amor, ni nadie meresçio
que ella lo estuuiesse. Que si otra
cossa entendiera, por tan dichosa
la tuuiera yo en morirse, como a
mí por desdichada en uer, o cruda
muerte, quan poco caso hazes de
mi: pues lleuandome todo mi bien,
me dexas, no para más, que para
sentir esta falta. O mi Arsileo, o
disçreçion jamás oyda, o el más
claro ingenio que naturaleza pudo
dar. ¿Qué ojos pudieron uerte,
qué animo pudo suffrir tu
desastrado fin? O Arsenio,
Arsenio, Arsenio quan poco
pudiste suffrir la muerte del
desastrado hijo, teniendo más
ocasion de suffrirla que yo? ¿Por
qué (cruel Arsenio) no quesiste
que yo partiçipasse de dos
muertes, que por estoruar la que
menos me dolia, diera yo çien mil
vidas, si tantas tuuiera? A Dios,
bienauenturada Nimpha, lustre y
honrra de la real casa de Aragon,
Dios dé gloria a tu anima, y saque
la mia de entre tantas
desuenturas. Despues Belisa vuo
dicho estas palabras, y despues
de auer uisto otras muchas
sepulturas, muy riquissimamente
labradas, salieron por una puerta
falsa que en el jardin estaua, al
verde prado: adonde hallaron a la
sabia Feliçia, que sola se andaua
recreando: la qual los reçibio con
muy buen semblante. Y en quanto
se hazia hora de çenar, se fueron
a vna gran alameda, que çerca de
alli estaua, lugar donde las
Nimphas del sumptuoso templo,
algunos dias salian a recrearse. Y
sentados en un pradezillo,
çercado de uerdes salzes,
començaron a hablar vnos con
otros: cada vno en la cosa que
más contento le daua. La sábia
Feliçia llamó junto a si al pastor
Sireno, y a Felismena. La Nimpha
Dorida, se puso con Syluano
hazia vna parte del verde prado, y
las dos pastoras, Seluagia, y
Belisa, con las más[1257]
hermosas Nimphas, Cinthia y
Polydora, se apartaron haçia otra
parte: de manera que aunque no
estauan vnos muy lexos de los
otros, podian muy bien hablar, sin
que estoruasse vno lo que el otro
dezia. Pues queriendo Sireno,
que la platica, y conuersaçion se
conformasse con el tiempo y
lugar, y tambien con la persona a
quien hablaua, començo a hablar
desta manera: No me paresçe