Yale Silviculture: Mark Ashton

	Download Research Statement (PDF, 376 KB) Publications Regeneration Studies Collaborations Silviculture Student Studies	View Slide Show Presentation of Research Research Sites: Meadow Lake, Saskatchewan PRORENA Sinharaja, Sri Lanka Yale Myers Forest, CT
	I foresee that upland forests in humid regions of the world, after two centuries of dramatic decline and degradation, will become critical resources for the sustenance of global services (water, climate amelioration, recreation) and products (genetic reservoirs of new products, specialty timber and non-timber products) demanded by society. My research concentrates on the ecological adaptations by which trees of various species of these complex forest types become established naturally after disturbances that make vacancies in the growing space. The kind of knowledge gained is a key part of the basis for developing silviculture that will sustain and augment the various forest values of the future.

The importance of big scale comparisons
The regeneration period is a critical window of time during which the future composition and development of the forest is largely determined. It is also the period during which the silviculturist has the most opportunity to restore and guide forest growth. My research has focused on understanding the most important biological and physical processes governing regeneration of species-rich moist forests. The majority of my research has been centered for over twenty years at field sites in two primary upland forest regions: 1) the American temperate moist deciduous forests (oak-hickory) of southern New England, USA (Yale Myers, Connecticut); and the Asian tropical wet-evergreen forest (mixed-dipterocarp) in Sri Lanka and Malaysia. For the last ten years I have also been working at three secondary field sites 1): the boreal spruce-aspen forests of Saskatchewan, Canada; 2) the moist mixed-conifer forests of interior British Columbia; and 3) the tropical seasonally-wet forests of Panama, Central America. Sites were selected to develop common methodological protocols that would enable a better understanding of the differences and similarities of regeneration within and across major forest biomes.

Long-term experimental rationale and framework: Building the basics
Cladistic groups (at the genus level in particular) are largely the level at which species differentiation occurs in tree species-rich forests such as those of eastern North America and the Asian moist tropics. Co-occurring species within a genus may differ in value and the products that they yield, as well as in their spatial and temporal role in biodiverse plant assemblages. I have chosen to study a series of co-occurring species that are of the same cladistic group (and often of the same successional status), because their similar morphology and growth adaptations facilitate examination of differences. Identifying these differences and their variations can provide a better understanding of evolving species specialization in relation to environment. This, in turn, provides the ecological information necessary for restoration and sustainable management of moist tree species-rich forests.

Applying this knowledge to silviculture
My work has immediate application for the development and testing of regeneration methods in natural forests, which I am pursuing at all my sites. I have long-term plots monitoring regeneration performance in experimental canopy openings that are intended to test hypotheses concerning forest resilience in relation to disturbance and site productivity. In the tropics I am using information gleaned from New England’s experience with deforestation, and together with my information on seedling regeneration ecology of the tropics (Panama, Sri Lanka), I have embarked on a series of sequential studies with collaborators on site reforestation. Much of this information has been summarized in two seminal textbooks on silviculture and agroforestry.

Collaboration
At all my research sites I have become part of a network of colleagues and interdisciplinary teams. I have a longstanding commitment to advising and nurturing young local scientists and practitioners. I have received several international prizes for my research, advising and teaching. My doctoral students have all followed similar themes and experimental protocols to my own for understanding regeneration dynamics of complex moist forests. Their studies, however, are based in other important forest regions of the Indian Himalayas, the mangrove deltas of Papua New Guinea, and the rain forests of Central Africa and the Peruvian Amazon. Lastly, teaching advanced-level graduate courses has provided me the opportunity to work with students and faculty on complex biological and social issues of forest use and protection. To date we have studied the Panama Canal watershed, the Venezuelan high Andes, community development in La Amistad Biosphere Reserve, Costa Rica, community forest management in Oaxaca, and watershed management issues of the Ecuadorian Andes.

Publications

Regeneration Studies       Silviculture         Collaborations      Student Studies

Comparative Regeneration Papers Published by Primary Core Research Sites

Sri Lanka         New England    Panama       British Columbia, Canada Reviews

Sri Lanka

Gunatilleke, C.V.S., Gunatilleke, I.A.U.N., Esufali, S., Harmes, K.E., Ashton, P.M.S., Burslem, D.F.R.P., & P.S. Ashton. ----. Species-habitat associations in a Sri Lankan dipterocarp forest. Journal of Tropical Ecology (in press).

Ashton, M.S., Singhakumara, B.M.P., & H.G. Gamage. 2006. Interaction between light and drought affect performance of tropical tree species that have differing topographic affinities. Forest Ecology and Management 207:110-122.

Tennakoon, S., Gunatilleke, I.A.U.N., Hafeel, K.M., Gunatilleke, C.V.S., & M.S. Ashton. 2005. Ectomycorrhizal colonization and seedling growth of Shorea (Dipterocarpaceae) species in simulated shade environments of a Sri Lankan rain forest. Forest Ecology and Management 205: 300-306.

Kathriarachchi, H.S., Tennakoon, K.U., Gunatilleke, C.V.S., Gunatilleke, I.A.U.N. & P.M.S. Ashton. 2004. Ecology of two selected liana species of utility value in a lowland rain forest of Sri Lanka: Implications for management. Conservation & Society 2: 273-288

Gunatilleke, C.V.S., Gunatilleke, I.A.U.N., Ethugala, A.U.K., Weerasekara, N.S., Ashton, P.S., Ashton, P.M.S., & S. Wijesundara. 2004 . Community ecology of a spatially explicit rain forest plot in Sinharaja, Sri Lanka. In "Forest diversity and dynamism: Results from the global network of large-scale demographic plots" Losos, E., & E. Leigh (Eds.). University of Chicago Press, Chicago, pp. 235-256.

Gamage, H.G., Singhakumara, B.M.P., & M.S. Ashton. 2004. Effects of light and fertilization on arbuscular mycorrhizal (AM) colonization and growth of tropical rain forest Syzygium tree seedlings. Journal of Tropical Ecology 20: 525-534.

Ashton, M.S., Gamage, H.K., & B.M.P. Singhakumara. 2003. Leaf structure of Syzygium Spp. (Myrtaceae) in relation to site affinity within a tropical rain forest. Botanical Journal of the Linnean Society 141: 365-377.

Singhakumara, B.M.P., Gamage, H.K., & M.S. Ashton. 2003. Comparative growth of four Syzygium species within simulated shade environments of a Sri Lankan rain forest. Forest Ecology and Management 174: 511-520.

Singhakumara, B.M.P., Uduporuwa, R.S.J.P., & P.M.S. Ashton. 2000. Soil seed banks in relation to light and topographic position of a hill dipterocarp forest. Biotropica 32: 190-196.

Gunatilleke, C.V.S., Gunatilleke, I.A.U.N., Ashton, P.M.S., & P.S. Ashton. 1998. Seedling growth of Shorea section Doona (Dipterocarpaceae) across an elevational range in southwestern Sri Lanka. Journal of Tropical Ecology 14: 231-245.

Gunatilleke, C.V.S., Gunatilleke, I.A.U.N., Perera, G.A.D., Burslem, D.F.R.P., Ashton, P.M.S., & P.S. Ashton. 1997. Responses to nutrient addition among seedlings of eight closely-related species of Shorea in Sri Lanka. Journal of Ecology 85: 301-311.

Cohen, A., Ashton, P.M.S., & B.M.P. Singhakumara. 1996. Releasing rainforest succession: A case study in the Dicranopteris linearis fernlands of Sri Lanka.   Restoration Ecology 3: 261-270.

Gunatilleke, C.V.S., Perera, G.A.D., Ashton, P.M.S., Ashton, P.S., & I.A.U.N. Gunatilleke. 1996. Seedling growth of Shorea section Doona, (Dipterocarpaceae) in soils from topographically different sites of Sinharaja rain forest in Sri Lanka. In "The Ecology of Tropical Forest Tree Seedling s". Swaine, M. D. (Ed.). pp. 245-266, UNESCO, Parthenon Press, Paris.

Ashton, P.M.S. 1995. Seedling growth of co-occuring   Shorea species in the simulated light environments of a rain forest. Forest Ecology and Management 72: 1-12.

Ashton, P.M.S., Gunatilleke, C.V.S., & I.A.U.N. Gunatilleke. 1995. Seedling survival and growth of four Shorea species in a Sri Lankan rainforest. Journal of Tropical Ecology 11: 263-279.

Ashton, P.M.S. 1992 . Some measurements of the microclimate within a Sri Lankan tropical rain forest. Agricultural and Forest Meteorology 59: 217-235.

Ashton, P.M.S., & G.P. Berlyn. 1992. Leaf adaptations of some Shorea species to sun and shade. New Phytologist 121: 587-596.

De Zoysa, N.D., & P.M.S. Ashton. 1991. The effect of different seed conditions, light and soil micro-environments on germination and survival of Shorea trapezifolia (Thw.) Ashton (Dipterocarpaceae). Jour. of Tropical Forest Science 3 (4):223-233.

Ashton, P.M.S., & N.D. de Zoysa. 1989. Performance of Shorea trapezifolia (Thw.) seedlings growing in different light regimes. Jour. of Tropical Forest Science 1 (4): 356-364.

New England, USA

Fladeland, M.M., Ashton, M.S., & X. Lee. 2003. Landscape variations in understory PAR for a mixed deciduous forest of southern New England. Journal of Agricultural and Forest Meteorology 118: 137-141.

Ashton, P.M.S., Yoon, K., Thadani, R., & G.P. Berlyn. 1999. Development of seedling leaf structure in relation to light environment of New England maples (Acer). Forest Science 45: 1-8.

Liptzin, D., & P.M.S. Ashton. 1999. Early successional stand dynamics of single-aged mixed hardwood stands in a southern New England forest, USA. Forest Ecology and Management.116: 141-150.

Ashton, P.M.S., Harris, P.G., & R. Thadani. 1998. Soil seed bank dynamics in relation to topographic position of a second-growth deciduous forest in southern New England, USA. Forest Ecology and Management 111: 15-22.

Li, W-L., Berlyn, G.P., & P.M.S. Ashton. 1996. Polyploids and their adaptation to water deficits in Betula papyrifera (Marsh). American Journal of Botany 83:15-21.

Ducey, M.J., Moser, W.K. & P.M.S. Ashton. 1996. Effect of fire intensity on understory composition and diversity in a laurel-dominated oak forest. Vegetatio 123: 81-90.

Ashton, P.M.S. & B.C. Larson. 1996. Germination and seedling growth of Quercus (section Erythrobalanus) across openings in a mixed-deciduous forest of southern New England, USA. Forest Ecology and Management 80: 81-94.

Moser, W.K., Ducey, M.J., & P.M.S. Ashton. 1996. Effects of fire intensity on competitive dynamics between red and black oaks and mountain laurel. Northern Journal of Applied Forestry . 13: 112-118.

Kittredge, D.B., & P.M.S.Ashton. 1995. Impacts of deer browse on regeneration in mixed-species stands in southern New England. Northern Jour. of Applied Forestry 12: 115-120.

Ashton, P.M.S. & G.P. Berlyn. 1994. A comparison of leaf physiology and anatomy of Quercus (section Erythrobalanus) species in different light environments. American Journal of Botany 81:589-597

Smith, D.M., & P.M.S. Ashton. 1993 . Early dominance of pioneer hardwoods after clearcutting and removal of advanced regeneration. Northern Jour. of Applied Forestry. 10(1):14-19.

Kittredge, D.B., & P.M.S.Ashton. 1990. Patterns of regeneration in mixed-species stands in southern New England. Northern Jour. of Applied Forestry 5:132-144.

Panama

Wishnie, M.H., Mariscal, E., Deago, J., Cedeño, N., Ibarra, D., Condit, R., Ashton, P.M.S., and D.H. Dent. ---- . Performance of 24 tropical tree species in relation to reforestation strategies in Panama. Forest Ecology and Management (in press).

Griscom, H.P., Kalko, E., & M.S. Ashton. ----- . Seed dispersers within a deforested, dry tropical region; Azuero Peninsula, Panama. Biotropica (in press).

Griscom, H.P., Ashton, M.S., & G.P. Berlyn. 2005. Seedling survival and growth of native tree species in pastures: implications for dry tropical forest rehabilitation in central Panama. Forest Ecology and Management 206:45-56.

British Columbia, Canada

Richardson, A.R., Berlyn, G.P., Ashton, P.M.S., McGroddy, M., & I.R. Cameron. 2001. Within-crown foliar plasticity of western hemlock, Tsuga heterophylla, in relation to stand age. Annals of Botany 56: 245-256.

Richardson, A.R., Berlyn, G.P., Ashton, P.M.S., Thadani, R., & I.R. Cameron. 2000. Foliar plasticity of hybrid spruce in relation to crown position and stand structure. Canadian Journal of Botany 78: 305-317.

Ashton, P.M.S.,Olander, L.P., Berlyn, G.P., Thadani, R., & I.R. Cameron. 1998. Changes in leaf structure in relation to crown position and tree size of Betula papyrifera within fire origin stands of interior cedar-hemlock, British Columbia. Canadian Journal of Botany 76: 1180-1187.

Reviews

Ashton, P.M.S. 1992. Establishment and early growth of advance regeneration of canopy trees in moist mixed-species broadleaf forest. In: "The Ecology and Silviculture of Mixed-Species Forests" Kelty, M.D., Larson B.C., & C.D. Oliver (Eds). pp. 101-125. Kluwer Academic Publ., Dordrecht, The Netherlands.

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Silviculture Studies

Studies     Reviews     Textbooks

Studies in New England, Sri Lanka, and Panama

Tomohiro, S., Ashton, M.S., Singhakumara, B.M.P., Gricom, H.P., Ediriweera, S. & B.W. Griscom. ---- . Effects of fire on the recruitment of rain forrest vegetation beneath Pinus caribaea plantations, Sri Lanka. Forest Ecology and Management (in press).

Ashton, P.M.S., Singhakumara, B.M.P., Evans, A.E., Gunatilleke C.V.S., and I.A.U.N. Gunatilleke. 2001 . A financial analyis of rain forest silviculture in southwestern Sri Lanka. Forest Ecology and Management 154: 431-444.

Ashton, P.M.S., Gamage, S., Gunatlleke, I.A.U.N., & C.V.S. Gunatilleke. 1998. Using Caribbean pine to establish mixed plantations: Testing effects of pine canopy removal on plantings of rain forest tree species. Forest Ecology and Management 106: 211-222.

Ashton, P.M.S., Samarasinghe, S.J., Gunatilleke, I.A.U.N., & C.V.S. Gunatilleke. 1997. Role of legumes in release of successionally arrested grasslands in the central hills, Sri Lanka. Restoration Ecology . 5: 36-43.

Ashton, P.M.S., Gamage, S., Gunatilleke, I.A.U.N., & C.V.S. Gunatilleke. 1997. Restoration of a Sri Lankan rain forest: Using Caribbean pine Pinus caribaea as a nurse for establishing late successional tree species. Journal of Applied Ecology 34: 915-925.

Samarasingha, J.N., Ashton, P.M.S., Gunatilleke, I.A.U.N., & C.V.S. Gunatilleke. 1996. Thinning guidelines for tree species of different successional status. Journal of Tropical Forest Science 10: 348-356.

Ashton, P.M.S., Gunatilleke, C.V.S., & I.A.U.N. Gunatilleke. 1993. A shelterwood method of regeneration for sustained timber production in Mesua-Shorea forest of southwest Sri Lanka. In : "Ecology and Landscape Management in Sri Lanka " Erdelen, W., Preu, C., Ishwaran, N., & M. Bandara (Eds.). pp. 255-274. Verlag Josef Margraf Scientific, Hamburg, Germany.

Ashton, P.M.S., Gunatilleke, I.A.U.N., & C.V.S. Gunatilleke. 1993. A Case for the development and evaluation of mixed-species plantations for the wet zone of Sri Lanka. In : "Ecology and Landscape Management in Sri Lanka" Erdelen, W., Preu, C., Ishwaran, N., & M. Bandara (Eds.). pp. 275-288. Verlag Josef Margraf Scientific, Hamburg, Germany.

Kittredge, D.B., Kelty, M.J. & P.M.S. Ashton. 1992 . The use of tree shelters with northern red oak in southern New England. Northern Jour. of Applied Forestry 9(4): 141-145.

Ashton, P.M.S., Lowe, J.S., & B.C. Larson. 1991 . Epicormic branching of blue mahoe (Hibiscus elatus L.). Jour. of Tropical Forest Science 3 (2):21-32

Ashton, P.M.S., Lowe, J.S., & B.C. Larson. 1990 . Thinning guidelines for blue mahoe (Hibiscus elatus L.). Jour. of Tropical Forest Science 2 (1): 37-47.

Ashton, P.M.S. 1990. A method for the evaluation of advanced regeneration for forest types of south and southeast Asia. Forest Ecology and Management 36:163-175.

Silviculture and Management Reviews

Ashton, M.S. 2003. The silviculture of dipterocarp forests in the Asian wet tropics. Forestry Chronicle 79: 263-267.

Ashton, P.M.S., Gunatilleke, C.V.S., Singhakumara, B.M.P., and I.A.U.N. Gunatilleke. 2001 . Restoration pathways for rain forest in southwest Sri Lanka: A review of concepts and models. Forest Ecology and Management 154: 409-430.

Ashton, M.S. , & C. Peters. 1999. Even-aged silviculture in mixed moist tropical forests with special reference to Asia: Lessons learned and myths perpetuated. Journal of Forestry 97: 14-19.

Ashton, P. M. S., & M. J. Ducey. 1997. The development of mixed plantations as successional analogs to natural forests. In "The National Proceedings: Forest and Conservation Nursery Associations 1996" Landis D., & D.B. South (eds.). U.S.D.A. Forest Service, Gen. Tech. Rep. PNW - 389.

Berlyn, G.P., & P.M.S. Ashton. 1997. Forests and the ecosystem paradigm. Journal of Sustainable Forestry 7: 141-157.

Berlyn, G.P., & P.M.S. Ashton. 1996 . Sustainability of forests. Journal of Sustainable Forestry . 3: 77-90.

Toman, M.A. & P.M.S. Ashton. 1996. Sustainable forest ecosystems and management: a review. Forest Science 42: 366-377.

Silviculture Textbooks

Ashton, M.S. & F. Montagnini (Editors). 1999. The Silvicultural Basis for Agroforestry Systems. CRC Press, Boca Raton, Florida. 278 p.
Smith, D.M., Larson, B.C., Kelty, M.J., & P.M.S. Ashton. 1997. The Practice of Silviculture: Applied Forest Ecology. 9th edition, Wiley & Sons, New York. 537 pp.

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Collaborative Publications

Clark, T.W., Cherney, D., Ashton, M.S., Bond, A., and A. Johnson. (in press). Journal of Sustainable Forestry, Special Issue 00: 000-000.

Clark, T.W., Dixon, L., Ashton, M.S., and B. Petit (Editors). 2006. Innovation and appraisal of sustainability efforts in La Amistad, Bocas del Toro, Panama and Talamanca, Costa Rica. Journal of Sustainable Forestry, Special Issue 23: 1-232.

Clark, T.W., Ashton, M.S., Zimmerman, K., and Q. Newcomer (Editors). 2004 . Conservation and development of the Condor Biosphere Reserve, Ecuador. Journal of Sustainable Forestry, Special Issue 18: 1-327.

Clark, T.W., Ashton, M.S. and J. Tuxill (Editors). 2002. Conservation or development? The case of the Amisconde project, La Amistad Biosphere Reserve, Costa Rica. Journal of Sustainable Forestry, Special Issue 16: 1-211.

Asbjornsen, H.A., & M.S. Ashton (Editors). 2001 . Balancing conservation with development: A comparison of two forest communities in Oaxaca, Mexico. Journal of Sustainable Forestry Special Issue 15: 1-131.

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Doctoral and Masters Student Studies

Brazil     Central Africa     India     Mexico     Peru     Malaysia

Brazil

Grogan, J., Landis, R.M., Ashton, M.S., & J. Galvao. 2004. Growth response by big-leaf mahogany (Swietenia macrophylla) advance seedling regeneration to overhead canopy release in southeast Para, Brazil. Forest Ecology and Management 204: 399-412.

Grogan, J., Ashton, M.S., & J. Galvao. 2003. Big-leaf mahogany (Swietenia macrophylla) seedling survival and growth across across a topographic gradient in southeast Para, Brazil. Forest Ecology and Management 186: 311-326.

Troy, A.R., Ashton, P.M.S., & B.C. Larson. 1997. Patterns in spatial distribution and abundance of a neotropical liana, Desmoncus polyacanthos, in relation to forest structure in the Brazilian Amazon. Economic Botany 51: 339-346.

Central Africa

Hall, J.S., McKenna, J.J., Ashton, P.M.S., & T.G. Gregoire. 2004. Habitat characterizations underestimate the role of edaphic factors controlling the distribution of Entandrophragma. Ecology: 85: 2171-2183.

Hall, J.S., Harris, D.J., Medjibe, V., and P.M.S. Ashton. 2003. The effects of selective logging on forest structure and tree species composition in a Central African forest: implications for management of conservation areas. Forest Ecology and Management 183: 249-264.

Hall, J.S., Medjibe, V., Berlyn, G.P., & P.M.S. Ashton.2003. Seedling growth of three co-occurring Entandrophragma species (Meliaceae) under simulated light environments: implications for forest management in Central Africa. Forest Ecology and Management 179: 135-144.

Hall, J.S., Ashton, M.S., and G.P. Berlyn. 2002. Seedling performance of sympatric Entandrophragma species (Meliaceae) under varied fertility and moisture regimes in a Central African rain forest. Journal of Tropical Ecology 18: 235-244.

NASA Remote Sensing Study on Landuse

Thenkabail, P.S., Enclona, E.A., Ashton, M.S., & Van Der Meer, V. 2005. Accuracy Assessments of Hyperspectral Waveband
Performance for Vegetation Analysis Applications. Remote Sensing of Environment 91: 1112-123.

Thenkabail, P.S., Stucky, N., Griscom, B.W., Ashton, M.S., Diels, J., Van Der Meer, B., & E. Enclona. 2004. Biomass estimations and
carbon stock calculations in the oil palm plantations of African derived savannas using IKONOS data. International Journal of Remote
Sensing 25: 5447-5472.

Thenkabail, P.S., Enclona, E.A., Ashton, M.S., Legg, C., Jean De Dieu, M. 2004. Hyperion, IKONOS, ALI, and ETM+ sensors in the
study of African rainforests. Remote Sensing of Environment 90: 23-40.

Thenkabail, P.S., Hall, J., Lin, T., Ashton, M.S., Harris, D. & E.A. Enclona. 2003. Detecting floristic structure and pattern across
topographic and moisture gradients in a mixed species Central African forest using IKONOS and Landsat-7 ETM+ images. International
Journal of Applied Earth Observation and Geoinformation 4: 255-270.

Himalaya, India

Quazi, S.A., Ashton, M.S., & R. Thadani. 2004. Regeneration of monodominant stands of banj oak (Quercus leuchotrichophora A. Camus) on abandoned terraces in the Central Himalayas. Journal of Sustainable Forestry 17: 75-90.

Thadani, R., & P.M.S. Ashton. 1995. Regeneration of banj oak (Quercus leucotricophora A. Camus) in the central Himalaya. Forest Ecology and Management 78: 217-224.

Mexico

Asbjornsen, H.A., Ashton, M.S., & D.J. Vogt. 2004. Effects of habitat fragmentation on the buffering capacityof edge environments in a seasonally dry tropical forest ecosystem in Oxaca, Mexico. Agricultural Ecosystems 103: 481-495.

Asbjornsen, H.A., Ashton, M.S., & K.V. Vogt. 2004. Synergistic response of oak, pine and shrub seedlings to edge environments and drought in a fragmented tropical oak forest, Oaxaca, Mexico. Forest Ecology and Management 192: 313-334.

Peru

Griscom, B.W., Daly, D., and P.M.S. Ashton. ----. Comparative floristics of lowland terra firme forests with and without dominance by aborescent bamboo (Guadua spp.) in southwestern Amazonia. Bulletin of the Torrey Bot. Club (in press).

Griscom, B., & M.S. Ashton. ---- . A self-perpetuating bamboo disturbance cycle in lowland terra firme forests of Madre de Dios, Peru. Journal of Tropical Ecology (in press).

Griscom, B.W., & M.S. Ashton. 2003. Bamboo control of forest succession: Guadua sarcocarpa Pilger in southeastern Peru. Forest Ecology and Management 175: 445-454.

Flores, C. F., & P.M.S. Ashton. 2000 . Harvesting impact and economic value of Geonoma deversa (Poiteau) Kunth, an understory palm used for roof thatching in the Peruvian Amazon. Economic Botany 54: 267-277.

Sarawak, Malaysia

Palmiotto, P.A., Davies, S.A., K. A. Vogt, P. M. S. Ashton, D.J. Vogt & P. S. Ashton. 2004. Soil related habitat specialization in dipterocarp raoin forest tree species in Borneo. Journal of Ecology 92: 609-623.

Palmiotto, P.A., Vogt, K.A., Ashton, P.M.S., Ashton, P.S., Vogt, D.J., Semui, H., & L.H. Seng. 2004 . Linking canopy gaps, topography and soils in a tropical rain forest: Implications for species diversity. In "Forest diversity and dynamism: Results from the global network of large-scale demographic plots" Losos, E., & R. Leigh (Eds.). University of Chicago Press, Chicago, pp 101-12.

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