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Propagating framework tree species to restore seasonally dry tropical

forest: Implications of seasonal seed dispersal and dormancy

Article in Forest Ecology and Management · July 2002

DOI: 10.1016/S0378-1127(01)00609-0

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 Forest Ecology and Management 164 (2002) 31–38

Propagating framework tree species to restore seasonally

dry tropical forest: implications of seasonal seed
dispersal and dormancy
David Blakesleya,*, Stephen Elliottb, Cherdsak Kuarakb, Puttipong Navakitbumrungb,
Sudarat Zangkumb, Vilaiwan Anusarnsunthornb
a
Horticulture Research International, East Malling, West Malling, Kent ME19 6BJ, UK
b
Science Faculty, Biology Department, Forest Restoration Research Unit, Chiang Mai University, Chiang Mai 50200, Thailand
Received 4 December 2000; received in revised form 10 April 2001; accepted 10 April 2001

Abstract

One effective approach to forest restoration in degraded tropical forestland is the so-called ‘framework species method’
which involves planting 20–30 indigenous forest tree species to re-establish a basic forest structure that catalyses the recovery
of biodiversity. For the seasonally dry tropical forests of Doi Suthep-Pui National Park in northern Thailand, a provisional list
of 36 potential framework species was compiled, from 19 different families representing a broad spectrum of the tree flora.
This paper examines the seed germination characteristics of these species when grown as a nursery ‘crop’ for planting to
restore degraded sites, focussing on germination phenology and dormancy. It considers how such characteristics affect the
first stage of nursery production from seed collection to pricking out seedlings in the nursery. Twenty-nine species had a
germination percentage of 60% or greater, which is acceptable for nursery production. The median length of dormancy (MLD)
ranged from 7 days in the case of Erythrina subumbrans to 219 days for Lithocarpus garrettianus. Germination was defined
as rapid if the MLD occurred within 3 weeks, and slow if occurring after 12 weeks. Twelve species germinated rapidly
and eight germinated slowly, the remainder being intermediate. Seedling emergence ranged over a period of 7 days in
the case of Erythrina stricta and E. subumbrans to 322 days in the case of L. garrettianus. # 2002 Elsevier Science B.V. All
rights reserved.

Keywords: Tropical forest restoration; Seed dormancy; Germination; Seedling development; Framework tree species

1. Introduction restore forests are increasing, but such efforts are

often limited due to lack of knowledge about how to
Deforestation is a serious environmental problem propagate the majority of indigenous tree species. One
throughout the tropics causing rural poverty, water- effective approach to forest restoration is the ‘frame-
shed degradation and loss of biodiversity. Efforts to work species method’ (Goosem and Tucker, 1995;
Lamb et al., 1997; Tucker and Murphy, 1997) first
developed to restore forest in degraded areas of
*
Corresponding author. Tel.: þ44-1732-84-3833;
Queensland’s Wet Tropics World Heritage Site in
fax: þ44-1732-84-9067. Australia. The method depends on tree planting to
E-mail address: [email protected] (D. Blakesley). restore basic forest structure which then encourages

0378-1127/02/$ – see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 1 ) 0 0 6 0 9 - 0
32 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38

the recovery of biodiversity. Seven years after planting work against the needs of small-scale tree nursery
20–30 framework tree species in degraded grassland managers.
sites in Queensland, the regenerating forests devel- A list of 36 potential framework species from 19
oped closed canopies up to 8.7 m tall and was different families was drawn up, based on pilot studies
naturally colonised by up to 49 additional tree species in the nursery, preliminary field trials over 3 years and
(Tucker and Murphy, 1997). Framework tree species fruiting characteristics. This included pioneers such as
are fast growing with dense spreading canopies which Melia toosendan and climax species such as H. dulcis.
rapidly shade out weeds. They also provide resources Key families include the Moraceae (four species),
for wildlife (such as fruit, nectar or perching sites) at Meliaceae (two species), Leguminosae (two species)
an early age. Animals (especially birds and bats) and Fagaceae (six species) (Table 1). The list is
attracted by such resources, disperse the seeds of necessarily provisional because long-term field trials
additional non-planted tree species into the planted are needed to determine the age at which the listed tree
sites, thus accelerating the return of biodiversity. Seed species first produce wildlife resources and the degree
of framework species should be easy to collect and to which they enhance biodiversity recovery. The
germinate in nurseries. A reasonable growth rate is present paper examines seed germination character-
also required in the nursery to ensure efficient use of istics of potential framework species when grown as a
nursery space and facilities. crop, focussing on dispersal, germination phenology
Although detailed information exists on the propa- and dormancy. It considers how such characteristics
gation of commercially valuable tree species, very affect the first stage of nursery production, from seed
little is known about potential framework tree species, collection to pricking out seedlings in the nursery. It
which tend to be non-commercial, indigenous forest also reviews the suitability of the species as frame-
tree species with high ecological value but low or work species based on the essential criterion of seed
unexplored economic value. For the vast majority of germination.
the huge diversity of forest tree species in Southeast
Asia flowering, fruiting and germination phenology
are not known and propagation techniques have not 2. Materials and methods
yet been developed. Of the 36 potential framework
species reported here, only Bishofia javanica, Dua- 2.1. Study site
banga grandiflora, Hovenia dulcis and Prunus
cerasoides have been studied previously (Datta and Trees were propagated in a nursery at 1000 m
Sharma, 1989; Frett, 1989; Kamaluddin and Grace, elevation near the headquarters of Doi Suthep-Pui
1993; Kopachon et al., 1996; Hardwick et al., 1997) National Park, north-west of Chiang Mai, northern
and none within the context of producing a ‘crop’ of Thailand (l88510 N latitude and 988540 E longitude).
framework species. The area experiences a monsoonal climate with
Producing a wide range of framework tree species is pronounced dry and wet seasons. Average annual
far more complex than mass propagation of a small precipitation recorded at nearby weather stations at
number of commercial plantation species. Indigenous similar elevations ranges from 1670 to 2094 mm. The
tree species in Thailand produce seeds at different wet season lasts from May to October and the dry
times throughout the year. However, seeds of tree season from November to April.
species in seasonally dry tropical forests in the neo- All the seed was collected in natural or slightly
tropics tend to germinate at the beginning of the rainy disturbed forest ecosystems close to the nursery
season (Garwood, 1983), providing seedlings with between elevations of 700–1600 m. This elevation
sufficient time to establish a good root system before range covers all the major forest types in the park,
onset of drought conditions during the following dry including the deciduous forest associations of the
season. However, it is not clear how germination lowlands (deciduous dipterocarp oak, bamboo decid-
phenology and year-round seed dispersal affect the uous forest and mixed evergreen deciduous forest) and
nursery operation. What may be the optimum strategy the evergreen forest of the uplands (Maxwell and
to enable trees to establish themselves naturally may Elliott, in press).
 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38 33

Table 1
Forest types, altitudinal ranges (northern Thailand) and fruit types of potential framework species

Species Family Forest typea Altitude range (m) Fruit type

Balakata baccata (Roxb.) Ess. Euphorbiaceae MED/E 400–500 Drupe

Bischofia javanica Bl. Euphorbiaceae BD/MED/E 525–1250 Drupe
Callicarpa arborea Roxb. var. arborea Verbenaceae DDO/BD/MED 375–1250 Berry
Castanopsis calathiformis (Skan) Rehd. and Wils. Fagaceae EP 1050–1500 Nut
Castanopsis tribuloides (Sm.) A. DC. Fagaceae MED/E/EP 900–1685 Nut
Cinnamomum iners Reinw. ex Bl. Lauraceae MED/E 700–1425 Berry
Debregeasia longifolia (Burm. f.) Wedd. Urticaceae MED/EP 525–1685 Achene
Duabanga grandiflora (Roxb. ex DC.) Walp. Sonneratiaceae MED/E 650–1450 Capsule
Eriobotrya bengalensis (Roxb.) Hk. f. forma Rosaceae E 1000–1650 Drupe
multinervata Vidal
Erythrina stricta Roxb. Leguminosae BD/E/EP 400–1680 Pod
Erythrina subumbrans (Hassk.) Merr. Leguminosae MED/E 500–1680 Pod
Eugenia albiflora Duth. ex Kurz Myrtaceae MED/E/EP 800–1525 Berry
Eurya acuminata DC. var. wallichiana Dyer Theaceae E 1000–1500 Berry
Ficus altissima Bl. Moraceae BD/MED 350–1050 Fig
Ficus racemosa var. racemosa Moraceae MED 350–500 Fig
Ficus semicordata B.-H. ex J.E. Sm. Var. semicordata Moraceae BD/E/EP 350–1550 Fig
Ficus subulata Bl. var. subulata Moraceae MED/E 825–1400 Fig
Gmelina arborea Roxb. Verbenaceae BD/MED/E/EP 350–1475 Drupe
Heynea trijuga Roxb. ex Sims Meliaceae BD/MED/E/EP 550–1680 Capsule
Hovenia dulcis Thunb. Rhamnaceae E 1025–1300 Capsule
Lithocarpus elegans (Bl.) Hatus. ex Soep. Fagaceae B/MED/EP 450–1450 Nut
Lithocarpus garrettianus (Craib) A. Camus Fagaceae B/MED/E 550–1100 Nut
Manglietia garrettii Craib Magnoliaceae E 1050–1600 Aggregate
follicle
Markhamia stipulata (Wall.) Bignoniaceae BD/MED/E/EP 950–1500 Capsule
Seem. ex K. Sch. var. kerrii Sprague
Melia toosendan Sieb. and Zucc. Meliaceae MED/E 700–1450 Drupe
Michelia baillonii Pierre Magnoliaceae MED/E 650–1100 Aggregate
follicle
Nyssa javanica Polygalaceae MED/E 550–1400 Drupe
Ostodes paniculata Bl. Euphorbiaceae E 1000–1350 Capsule
Phoebe lanceolata (Nees) Nees Lauraceae MED/E/EP 550–1550 Drupe
Planchonell punctata Flet. Sapotaceae DDO/BD/MED/E/EP 350–1525 Berry
Prunus cerasoides D.Don Rosaceae MED/E/EP 1050–1685 Drupe
Quercus semiserrata Roxb. Fagaceae MED/E/EP 800–1675 Nut
Quercus vestita Rehd. and Wils. Fagaceae E/EP 1200–1600 Nut
Rhus rhetsoides Craib Anacardiaceae MED/E/EP 650–1550 Drupe
Sapindus rarak DC. Sapindaceae MED/E 625–1620 Drupe
Spondias axillaris Roxb. Anacardiaceae MED/E/EP 700–1600 Drupe
a
BD: bamboo and deciduous; DDO: deciduous dipterocarp oak; MED: mixed evergreen and deciduous; E: evergreen; EP: evergreen and
pine (sensu Maxwell and Elliott, in press).

2.2. Seed germination were ‘fresh’ and undecayed. Following the removal of
the fruit pericarp, seeds were sown within 2–3 days
Seeds of the 36 potential framework species were of collection into modular plastic trays, on to the
collected from single parent trees of each species surface of a medium of two parts forest soil to one
when fruits were mature and ripe. Fruits were cut from part coconut husk. For each species, 72 seeds were
branches or collected from the ground only if they divided into three replicate batches of 24 which were
34 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38

randomly assigned to different benches and watered partially shaded under a transparent plastic roof
daily. Each replicate consisted of 24 adjacent compar- (approximately 40% full sunlight, similar to the light
tments ð3:5 cm  3:0 cm  7:0 cmÞ in one-seed tray. intensity in partially regenerating gaps). Once the first
Seed trays were placed on the top of concrete benches, pair of leaves had fully expanded, seedlings were

Table 2
Seed germination data of potential framework tree species, suitable for forest restoration plantings in northern Thailand

Species Seed collection Mean germination MLD (days)b Time over which Germination and
month percentagea (S.D.) seeds germinated synchrony
(days)b categoriesc

Balakata baccata (Roxb.) Ess. December 25 (6.3) 67 112 IG/AS

Bischofia javanica Bl. November 43 (14.6) 85 154 SG/AS
Callicarpa arborea Roxb. var. arborea August 67 (21.7) 86 63 SG/IS
Castanopsis calathiformis (Skan) June 61 (19.2) 16 42 RG/IS
Rehd. and Wils.
Castanopsis tribuloides (Sm.) A. DC. September 83 (8.3) 31 42 IG/IS
Cinnamomum iners Reinw. ex Bl. April 75 (8.3) 17 63 RG/IS
Debregeasia longifolia (Burm. f.) Wedd. March 100 (0) 15 14 RG/S
Duabanga grandiflora (Roxb. ex DC.) Walp. April 86 (2.9) 31 42 IG/IS
Eriobotrya bengalensis (Roxb.) September 79 (3.6) 16 203 RG/AS
Hk. f. forma multinervata Vidal
Erythrina stricta Roxb. May 67 (33.3) 10 7 RG/S
Erythrina subumbrans (Hassk.) Merr. April 39 (2.4) 7 7 RG/S
Eugenia albiflora Duth. ex Kurz May 71 (12.5) 24 147 IG/AS
Eurya acuminata DC. var. wallichiana Dyer March 69 (6.4) 60 126 IG/AS
Ficus altissima Bl. March 97 (2.4) 34 105 IG/AS
Ficus racemosa var. racemosa February 92 (4.2) 27 70 IG/IS
Ficus semicordata B.-H. ex J.E. March 92 (5.0) 52 41 IG/IS
Sm. var. semicordata
Ficus subulata Bl. var. subulata January 71 (8.3) 60 175 IG/AS
Gmelina arborea Roxb. March 83 (18.2) 25 14 IG/S
Heynea trijuga Roxb. ex Sims November 83 (14.4) 96 203 SG/AS
Hovenia dulcis Thunb. August 71 (15.0) 97 154 SG/AS
Lithocarpus elegans (Bl.) Hatus. ex Soep. September 69 (9.6) 143 231 SG/AS
Lithocarpus garrettianus (Craib) A. Camus September 56 (37.3) 219 322 SG/AS
Manglietia garrettii Craib October 74 (4.8) 81 140 IG/AS
Markhamia stipulata (Wall.) Seem. ex K. March 56 (2.1) 13 15 RG/.S
Sch. var. kerrii Sprague
Melia toosendan Sieb. and Zucc. April 67 (15.0) 15 70 RG/IS
Michelia baillonii Pierre June 31 (6.9) 101 63 SG/IS
Nyssa javanica July 67 (19.1) 39 70 IG/IS
Ostodes paniculata Bl. November 53 (16.8) 124 203 SG/AS
Phoebe lanceolata (Nees) Nees April 79 (4.2) 44 56 IG/IS
Planchonella punctata Flet. June 89 (1.7) 17 35 RG/IS
Prunus cerasoides D.Don March 74 (4.8) 52 63 IG/IS
Quercus semiserrata Roxb. June 92 (7.2) 18 35 RG/IS
Quercus vestita Rehd. and Wils. September 74 (13.4) 14 21 RG/S
Rhus rhetsoides Craib December 50 (50.0) 24 28 IG/IS
Sapindus rarak DC. January 83 (8.3) 45 98 IG/AS
Spondias axillaris Roxb. March 43 (4.8) 11 21 RG/S
a
Three replicates.
b
Pooled replicates.
c
RG: rapid germination; IG: intermediate germination; SG: slow germination; S: synchronous; IS: intermediate synchrony; AS:
asynchronous.
 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38 35

pricked out and transplanted into individual contain- and slow if the MLD was 84 days or more. Twelve
ers. Germination was monitored throughout the ger- species could be classified as having rapid germina-
mination period and was defined as emergence of tion: Castanopsis calathiformis; Cinnamomum iners;
any part of the shoot. The dates of the first and last Debregeasia longifolia; Eriobotrya bengalensis; Ery-
seeds to germinate were recorded, and the median thrina stricta; Erythrina subumbrans; Markhamia
length of dormancy (MLD) calculated (pooling stipulata; Melia toosendan; Planchonella punctata;
individuals of each species from the three replicate Quercus semiserrata; Quercus vestita; Spondias
batches) from the germination times of all seeds which axillaris. In contrast, Bischofia javanica, Callicarpa
germinated. arborea, Heynea trijuga, Hovenia dulcis, Lithocarpus
elegans, Lithocarpus garrettianus, Michelia baillonii
and Ostodes paniculata were categorised as having
3. Results slow germination. The remaining 16 species had
MLD’s of between 3 and 12 weeks and could be
Germination percentage, one of the key selection regarded as having intermediate germination rates.
criteria for framework species, ranged from 25 to Considering the framework species as a whole,
100% (Table 2). However, 80% of species had a most species (28 or 78%) fell into the categories of
germination percentage of 60% or greater, which is rapid or intermediate germination. Of the 21 species
more acceptable for this type of nursery operation. collected in the late dry and early wet season, only one
Only three species had low germination percentages: species, Michelia baillonii germinated slowly (Fig. 1).
Balakata baccata (25%); Michelia baillonii (31%); In contrast, of the 15 species collected in the late wet
Erythrina subumbrans (39%). However, these species and early dry season, seven species germinated slowly
still qualify as potential framework species due to (19% of the total); the remaining eight were inter-
other attributes, such as high growth rate in containers mediate or rapid. This seasonal variation resulted in a
or good field performance (unpublished data). peak in nursery germination in the first-half of the year,
The MLD ranged from 7 to 219 days. For the when the median seeds of 72% of species germinated
purposes of nursery production, germination was (Fig. 2). This coincided with the end of the latter part of
defined as rapid if the MLD was 21 days or less, the dry season and the early part of the wet season.

Fig. 1. The relationship between the MLD and the month of seed collection of species collected in Doi Suthep-Pui National Park (700–
1600 m asl). Each point represents an individual species.
36 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38

Fig. 2. Number of species located in Doi Suthep-Pui National Park (700–1600 m asl), whose median seed emergence falls in each month.

Seedling emergence ranged over 7 days for both Hardwick et al. (1997) studied germination and
Erythrina spp. to 322 days for L. garrettianus. For tree emergence of Prunus cerasoides collected on Doi
production in the nursery, germination was defined as Suthep, and also found that it fruited late in the dry
synchronous if all seedlings of a given species season with a high germination percentage. Bischofia
emerged within 21 days, and highly asynchronous if javanica has previously been propagated from seed,
this occurred over a period of more than 84 days. and grown in controlled environments to stimulate
Seven species germinated synchronously, six of which different forest canopies (Kamaluddin and Grace,
also had an MLD of less than 21 days (Table 2): 1993). This study showed that B. javanica has a wide
Debregeasia longifolia; Erythrina stricta; Erythrina acclimation potential to the changing light levels, which
subumbrans; Markhamia stipulata; Quercus vestita; may occur in gaps. There is a report of soft rot on
Spondius axillaris. The other species which germi- seedlings of Duabanga grandiflora (Datta and Sharma,
nated synchronously, Gmelina arborea, also germi- 1989). Although there are a number of other publi-
nated relatively rapidly, with an MLD of 25 days. cations relating to related taxa within the families
Species exhibiting highly asynchronous germination reported here, particularly in America and the neo-
were distributed across intermediate- and slow- tropics, no other relevant work has been published
germinating species. Of the latter group of species, on the potential framework species described in this
none germinated synchronously; 63 days was the paper.
shortest time of seedling emergence, and the mean Because of the rainfall patterns in a seasonally dry
emergence time for the eight species was 164 days. tropical forest, the ideal time to plant out container-
grown tree seedlings is at the start of the wet season. It
is a considerable challenge to produce a crop of
4. Discussion seedlings, of a plantable size, of 36 framework tree
species, all to be dispatched at the same time of year
Few phenological studies have been reported with when seeds are available at different months through-
the framework species described in this paper. The most out the year and they exhibit widely different rates of
studied has been Hovenia dulcis, with several reports on germination and growth in the nursery. The present
seed germination (Frett, 1988, 1989; Kopachon et al., study has shown that nursery production of such a
1996) and the successful micropropagation of axillary ‘collection’ of native species, about which very little is
buds from mature trees (Echeverrigaray et al., 1998). known, presents considerable logistical problems for
 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38 37

the nursery manager, even to get the seedlings to the research described in this paper. Other donors have
point of pricking out into containers. The first of these included The Fagus Anstruther Memorial Trust, The
is that to propagate 36 framework species, at least one Peter Nathan Trust, The Robert Kiln Charitable Trust,
collection trip would be required in every month of the The Barbara Everard Trust for Orchid Conservation,
year and probably more in March, April and Mr. Alan and Mrs. Thelma Kindred, Mr. Nostha
September when 18 species (50% of the framework Chartikavanij, Mr. R. Butterworth and Mr. James C.
species) are available for collection. Furthermore, Boudreau. The authors thank J.F. Maxwell for iden-
these species exhibit considerable variation in dor- tifying the tree species named in this paper. Voucher
mancy and germination synchrony. It can be clearly specimens are stored at the Chiang Mai University
seen from the scatter plot of MLD (Fig. 1) that species Herbarium, Biology Department. The authors are
with seeds dispersed in the late dry/early wet season grateful to all research assistants and volunteers who
tend to germinate quickly in the nursery, whereas assisted with data collection and processing and care
those with species dispersed towards the end of the wet of the plants in the nursery including: Jumpee
season and into the dry season, are likely to have a Bunyadit, Thonglaw Seethong, Tim Rayden, Kevin
much longer dormancy period. Seven of the frame- Woods, Rungtiwa Bunyayod and Janice Kerby. The
work species appear to be ideal for nursery production, Head and staff of Doi Suthep-Pui National Park
because they are collected at one time of the year in the Headquarters provide essential collaboration for
late dry/early wet season (with the exception of FORRU’s research. We are especially grateful to
Quercus vesita) and germinate rapidly and synchro- the Head of the National Park, Mr. Paiboon Sawet-
nously. These species, therefore, require minimum melanon, Mr. Amporn Panmongkol (Deputy Head)
time in the germination facility where they are and Mr. Prasert Saentaam.
particularly susceptible to pests and diseases. The
other species collected at this time (with the exception
References
of Michelia baillonii) have intermediate germination,
and vary in the synchronicity of germination, and
Datta, B., Sharma, G.D., 1989. Report on soft rot of Duabanga
include species such as Eurya acuminata and Ficus grandiflora seedling (Roxb. ex DC.) Walp. Curr. Sci. 58, 574–
subulata which are highly asynchronous. Another 575.
predictable group of species, in terms of nursery Echeverrigaray, S., Mossi, A.J., Munari, F., 1998. Micropropaga-
planning, are those dispersed in the late wet/early dry tion of raisin tree (Hovenia dulcis Thunb.) through axillary bud
culture. J. Plant Biochem. Biotechnol. 7, 99–101.
season which germinate slowly and also asynchro-
Frett, J.J., 1988. Requirements for germination of Hovenia dulcis
nously. seeds. Hortscience 23, 677.
Whilst it is beyond the scope of this paper to Frett, J.J., 1989. Germination requirements of Hovenia dulcis
consider the manipulation of growth and development seeds. Hortscience 24, 152.
of framework species in containers, it is clear from the Garwood, N.C., 1983. Seed germination in a seasonal tropical
forest in Panama: a community study. Ecol. Monogr. 53, 159–
above discussion that seedlings will be ready for
181.
pricking out throughout the year. Further work is now Goosem, S.P., Tucker, N.I.J., 1995. Repairing the Rainforest—
underway to assess the second stage of nursery Theory and Practice of Rainforest Re-establishment in North
production of these species, from pricking out through Queensland’s Wet Tropics. Wet Tropics Management Author-
to weaning and dispatch. ity, Cairns, pp. 71.
Hardwick, K., Healey, J., Elliott, S., Garwood, N.C., Anusarnsun-
thorn, V., 1997. Understanding and assisting natural regenera-
tion processes in degraded seasonal evergreen forests in
Acknowledgements northern Thailand. For. Ecol. Manage. 99, 203–214.
Kamaluddin, M., Grace, J., 1993. Growth and photosynthesis of
FORRU was funded with financial support from tropical forest tree seedlings (Bischofia javanica Blume) as
influenced by a change in light availability. Tree Physiol. 13,
Riche Monde (Bangkok) Ltd. and United Distillers
189–201.
PLC. Shell Forestry Limited, the Biodiversity Kopachon, S., Suriya, K., Hardwick, K., Pakaad, G., Maxwell, J.F.,
Research and Training Programme and the Science Anusarnsunthorn, V., Garwood, N.C., Blakesley, D., Elliott, S.,
Faculty of Chiang Mai University sponsored the 1996. Forest restoration research in northern Thailand. 1.
38 D. Blakesley et al. / Forest Ecology and Management 164 (2002) 31–38

Fruits, seeds and seedlings of Hovenia dulcis Thunb. Nat. Hist. Maxwell, J.F., Elliott, S., in press. Vegetation and Vascular Flora
Bull. Siam. Soc. 44, 41–52. of Doi Suthep-Pui National Park, Chiang Mai Province,
Lamb, D., Parrotta, J., Keenan, R., Tucker, N.I.J., 1997. Rejoining Thailand. The Biodiversity Research and Training Programme,
habitat remnants: restoring degraded rainforest lands. In: Bangkok.
Laurence, W.F., Bierrgaard Jr., R.O. (Eds.), Tropical Forest Tucker, N.I.J., Murphy, T.M., 1997. The effects of ecological
Remnants: Ecology, Management and Conservation of Frag- rehabilitation on vegetation recruitment: some observation from
mented Communities. University of Chicago Press, Chicago, the Wet Tropics of North Queensland. For. Ecol. Manage. 99,
IL, pp. 366–385. 133–152.

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