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Plant species’ attributes & spatial patterns of regeneration in secondary rainforests – combining perspectives from Spatial Ecology and Plant Functional Ecology Overview In view of extensive loss, fragmentation and disturbance of rainforests over the past century, conservation of rainforest species and ecosystems depends not only on protection of remnant rainforests, but on understanding the ecology of ‘secondary’ rainforests and their regeneration. This project quantifies spatial patterns of plant regeneration in primary and secondary rainforests, and relates the observed distributions to : Jessie Wells, PhD Student i) spatial distributions of rainforest cover and source plants [email protected] ii) ecological traits of plant species - dispersal mechanisms, age of reproduction, plant life-span, & attributes of seeds, stems, and leaves. The Ecology Centre University of Queensland Research Plans Past fenceline The project’s two components combine studies of plant species attributes with spatial analyses of regeneration in secondary rainforests on abandoned pastures, and in adjacent, intact rainforests. Comprehensive transect studies Transect 1 Transect 2 Two transects 180 x 10 m – from primary into secondary rainforest – all plants above 4th-leaf seedlings 1) Comprehensive studies Changes in species composition, diversity, and the representation of plant functional traits along transects from primary into secondary rainforest Year 1964 – to study variation over distance from the primary forest, in strata from seedlings to the canopy – considering six ‘principal traits’, and quantifying representation as % of species and % of stems. Comprehensive site Key Questions – Comprehensive studies Atherton Tablelands,Wet Tropics – rainfall > 4000 mm p.a, basalt soils, 390 m a.s.l. 1) Plant functional traits - inter-relations and spatial variation What are the main dimensions of variation between species ? – to what extent are traits co-ordinated to form alternative sets or strategies ? Are there changes in the diversity or relative representation of functional traits with distance from primary forest, and between height classes of regeneration ? – for example: declines in the diversity of seed sizes present, or declines in the % of species with specific dispersers such as musky rat-kangaroos. 2) Species composition and diversity Predicted changes in diversity closer to the primary rainforest: i) higher richness, lower dominance & higher rarity, ii) increased inter-dispersion of species and of size-classes – reflecting a temporal shift from regeneration of several light-demanding species, to more probabilistic and continuous regeneration of many other species. – analysis of a more-extensive set of functional traits and their inter-relations – spatial modelling of regeneration for individual species, developing statistical ‘summed regeneration shadow’ models for a range of regeneration classes. Set 1 – Wet Tropics - 21 tree & shrub species, Robson Creek Set 2 – Wet Tropics CSIRO - 13 large-seeded trees, 4 fragments, 4 primary sites Set 3 – La Selva, Costa Rica – traits of 11 species, mapping of 1 subcanopy palm – examples from Site 2, 40 yrs regeneration Sapindaceae 0.8 to 1.6m 450 The diversity seen in any given aspect of plant form and function can be extensive - however, this does not lead to an infinite diversity of combinations, due to trade-offs and interdependencies among traits. Key questions in plant functional ecology concern the inter-relations of observable plant features, and how they reflect aspects of ecological behaviour and evolution. An objective is to quantify traits that give the most direct indication of ecological mechanisms of dispersal, establishment and persistence that are central to the regeneration of any plant, and form the basis of plant strategy theories. Principal plant traits Traits reflecting alternative resource allocations to growth & activity versus persistence, resource efficiency & conservation: 4) Leaf traits – Specific Leaf Area, Water content, Leaf Strength 5) Maximum Height 6) Stem tissue Density Further traits considered in focal species studies: lifespan, growth rates, mortality, fruiting phenology. Key Questions – Focal Species studies Diploglottis bracteata > 1.6m tall Secondary forest age 40 yrs Primary forest Plant functional traits 1) Seed mass 2) Reproductive Age & Light environment 3) Dispersal mechanisms – wind, bats, rats, fruit pigeons, cassowary… 2) Focal Species Plant distributions Year 2000 7863 plants 196 species 67 families – large seeds with aril – dispersed by cassowary & musky rat kangaroo – rainforest subcanopy tree seedlings 0.2 to 0.8m 440 Adults 430 1) Point patterns – Conspecific aggregation / dispersion, relative neighbourhood density statistics 2) Spatial signatures of regeneration –‘Summed regeneration shadow’ models – based on the distribution of possible source-plants, and analysis of observed regeneration to estimate seed-source strengths, and probability functions for dispersal distances, using Likelihood methods. The regeneration shadow’s ‘intensity’ at a given point in space is a summed contribution from each possible source-plant, depending on its source strength and distance. 420 410 400 390 170 180 190 200 210 220 230 240 250 260 Ceratopetalum succirubrum Cunoniaceae – large winged seeds – ‘flutter’ dispersal by wind – canopy tree of uplandmountain rainforests 270 280 290 300 Regeneration Shadow = Σ Source strength x Distance function Secondary forest age 40 yrs Primary forest Distance-function examples 450 440 density (m-2) 160 430 420 > 1.6m tall 410 0.8 to 1.6m seedlings 0.2 to 0.8m 400 Adults 390 160 170 180 190 200 210 220 230 240 250 260 270 280 290 300 distance (m) Secondary forest age 40 yrs Primary forest Cryptocarya mackinnoniana Lauraceae – medium size seeds – dispersed by a diverse set of bird species – subcanopy tree, many reaching maturity 450 440 430 420 410 Aerial view - distribution of sources (circles, scaled by tree diameters) and contours of seed density (Clark et al.1999, fig 2 p.1479) distance (m) Observed patterns of regeneration result from processes of seed-production, dispersal, and seed & seedling survival, and I cannot analyse these transitions directly. However, models of 2-3 seedling stages should indicate the importance of source distributions & seed dispersal for species with contrasting dispersal mechanisms and plant attributes. 400 390 160 170 180 190 200 210 220 230 240 250 260 270 280 290 300 For animal-dispersed species, the relative roles of dispersal, predation, and seedling survival will be examined by comparing results from i) empirical models of observed regeneration, and ii) process models of frugivory, seed deposition & seed predation (D.Westcott & A.Dennis, CSIRO). Contributions Ecological Theory Restoration Ecology – how quantifiable attributes and the spatial patterns of plants relate to concepts of their functional ecology & regeneration strategies – possibilities & limitations of natural regeneration for rainforest conservation – how management actions directed to limiting factors such as seed dispersal could assist the regeneration of species & functional diversity.