What determines plant species invasiveness? A case-study of Amani Botanical Garden, Tanzania

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1 What determines plant species invasiveness? A case-study of Amani Botanical Garden, Tanzania Wayne Dawson 1, Phillip E. Hulme 2 and David F.R.P. Burslem 3 1 Centre for Ecology and Hydrology/ University of Aberdeen, UK 2 National Centre for Advanced Bio-Protection Technologies, NZ 3 University of Aberdeen, UK

2 What determines invasion success? Problems with trait-based analyses -Many confounding variables Need to control for -Recipient ecosystem -Residence time -Introduction effort Botanical Gardens may offer opportunities

3 Amani History Established in 1902 by Germans -Continued management by British after WW1; -Management ceased after 1970s Detailed records and maps of plantations and species planted -601 species introduced between 1902 and Total area 350 ha. 20 plantations subdivided into 141 compartments, containing 1978 species plots (Greenway, 1934) Honess (1963) ~400 spp remaining in 1005 plots

4 Amani Geography ABG embedded within Amani Nature Reserve, East Usambara mountains, NE Tanzania Part of Eastern Arc Mountains (biodiversity hotspot) ANR conserves submontane and lowland rainforest (8000ha.) - high conservation value A number of species now recognised as at least locally invasive and threaten forest 3 km

5 Assessing invasion status of species from ABG collections Initial baseline survey of ABG plantations -Revisited all 1005 species plots surviving in Recorded species survival + presence of other unplanted alien species Opportunistic forest survey -Forest fragments and edges adjacent to plantations

6 Assessing invasion status of species from ABG collections Classifying species invasion status- -Surviving only: Original adults present, but no evidence of regeneration -Regenerating only: Seedling and saplings present, but no new adults -Naturalising only: New adults present, but spreading only locally (<10 new compartments) -Spreading: Naturalised, and spreading considerably (>10 new compartments). -Found in forest

7 Of 211 species surviving (out of 601) 138 species are surviving only - Eucalyptus spp.

8 Of 211 species surviving (out of 601) 138 species are surviving only - Eucalyptus spp. 35 species are regenerating only - Ceiba pentandra

9 Of 211 species surviving (out of 601) 138 species are surviving only - Eucalyptus spp. 35 species are regenerating only - Ceiba pentandra 32 species are only naturalised - Artocarpus heterophyllus

10 Of 211 species surviving (out of 601) 138 species are surviving only - Eucalyptus spp. 35 species are regenerating only - Ceiba pentandra 32 species are only naturalised - Artocarpus heterophyllus 17 species are spreading - Syzygium jambos

11 Of 211 species surviving (out of 601) 138 species are surviving only - Eucalyptus spp. 35 species are regenerating only - Ceiba pentandra 32 species are only naturalised - Artocarpus heterophyllus 17 species are spreading - Syzygium jambos 25 species found in forest - Castilla elastica

12 Spreading species in Amani are naturalised/ invasive elsewhere 1-PIER species list 2-Binggeli (1998) Overview of invasive plants in the Tropics 3-Haysom & Murphy (2003) Invasive Forestry Trees 25 2 =14, df=5, p=0.01 Naturalised/invasive elsewhere Species frequency Not naturalised/invasive elsewhere 0 Regenerating Naturalising Spreading Invasion status

13 Invasion status and rate of spread All spreading species with known time since introduction have spread > 300m in 50 years One naturaliser has spread >>300m- Coffea canephora Spread in 50 years (m) Regen Naturalised Spreading

14 Greater effort Introduction effort and invasiveness Demographic/ environmental stochasticity Greater probability of survival and naturalisation Surviving species planted in greater numbers than species lost log(number of plants) Absent Surviving Regen Naturalised Spreading

15 What determines invasiveness? Climate and latitude -Climate similar in native range to introduced range -Greater survival rates, growth and reproduction Tropical species (equatorial, wet) expected to be more successful Amani

16 Native climate and invasiveness Species only surviving from higher latitudes 25% of survivors only are Eucalyptoid (14spp) and 23% are gymnosperms (13spp) Midpoint of latitudinal range (degrees) Surviving Regenerating Naturalised Spreading

17 What determines invasiveness? Species traits Small seed size Greater seed output Fast germination Fast Growth Invaders of disturbed areas Dispersal -Wind dispersal successful in open habitats -Dispersal by tree dwelling animals (birds, primates, squirrels) successful in closed semi-natural/natural vegetation

18 Seed size Spreading species have marginally smaller dry seed weights than regenerating and naturalised species But so do survivors Who fail to regenerate! log10(dry seed weight, mg) Surviving Regenerating Naturalised Spreading

19 Herbivory and invasiveness Enemy Release- Species encountering fewer enemies in introduced range have an advantage Proportion of leaves damaged correlated with no. native confamilial species (rho=0.47, p=0.01) % Area leaf damage not correlated with no. native confamilial species (rho=0.16, p=0.41) Proportion of leaves damaged Number of native confamilial species

20 Herbivory and invasiveness Spreading and non-spreading spp. do not differ in proportion of leaves damaged (t=-0.926, df=24 p=0.364) Proportion of leaves damaged Not spreading Spreading

21 Current and further work Compile more data on species traits and ecological variables -Shade Tolerance -Growth (slow, moderate, fast) -Native Vegetation type (dry, moist) -Temperature and rainfall Look at relative roles of all factors in explaining species success at different stages of invasion Investigate potential importance of leaf traits (specific leaf area and nutrient content) Is invasiveness related to herbivory (enemy release)-control for family Test effectiveness of Hawaiian Weed Risk Assessment in identifying high and low risk species

22 Thanks to- Asante sana kwa kuni sikilize! Natural Environment Research Council/ Defra Darwin Initiative Tropical Biology Association (TBA) especially Dr. Rosie Trevelyan All staff at Amani Nature Reserve, especially Mr C.T. Sawe (Conservator) Botanists Ahmed S Mndolwa and Iddy Rajabu (Tanzania Forestry Research Institute) To find out more about the Darwin Initiative Project Combating Invasive Alien Plants Threatening The East Usambara Mountains, Tanzania visit:

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