SOME CHARACTERISTICS of a NATIVE ENVIRONMENTAL WEED: PITTOSPORUM UNDULATUM Trudi L
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Twelfth Australian Weeds Conference SOME CHARACTERISTICS OF A NATIVE ENVIRONMENTAL WEED: PITTOSPORUM UNDULATUM Trudi L. Mullett School of Ecology and Environment, Deakin University, 662 Blackburn Road, Clayton, Vic. 3168 (Current Address: PO Box 160, Murchison, Vic. 3610) Abstract Invading populations of Pittosporum features that make invasive populations of P. undulatum undulatum Vent. (Sweet Pittosporum) impact on the so difficult to manage in invaded remnant vegetation. floristic and structural composition of invaded plant The abundant winter fruit crops produced by ornamen- communities. This plastic, adaptable species is able to tal plantings and established populations of P. colonise a range of environments and is arguably the undulatum attract a suite of generalist avian frugivores most successful native weed in south east Australia. including Pied Currawongs (Strepera graculina), Eu- Abundant P. undulatum fruit crops attract a suite of ropean Blackbirds (Turdus merula), Satin Bowerbirds generalist, opportunistic frugivores and seed (Ptilonrhynchus violaceus) Silvereyes (Zosterops lat- germinability is high in natural and invaded habitats. eralis) and Red whiskered Bulbuls (Pycnonotus The continued invasion of P. undulatum in remnant jocosus) (Cooper 1959, Gleadow 1982, Emison et al. vegetation is further facilitated by fundamental changes 1987, Buchanan 1989, Barker and Vestjans 1990, in natural fire regimes. Mullett 1996). Impacts associated with P. undulatum invasion vary in The establishment of P. undulatum individuals is gen- different habitats and are most obvious in dry erally restricted to the base of mature trees as these sclerophyll forest. Changes in the composition of in- offer perching sites for frugivorous dispersers, which digenous species are evident with incremental increases defecate or regurgitate undigested seeds whilst perch- in P. undulatum cover-abundance. Considerable de- ing. The clumps of P. undulatum which develop act as clines in indigenous species richness have been re- a focus for further avian activity and thus for further corded at some invaded sites. The floristic and struc- seed dispersal (Mullett and Simmons 1995). The proc- tural simplification associated with P. undulatum in- ess of clump enlargement perpetuates and as these vasion reduces the diversity of habitat resources and clumps coalesce virtual monospecific stands of P. ultimately diminishes the biodiversity values of invaded undulatum develop (Richardson and Brink 1985, sites. Mullett 1996). INTRODUCTION P. undulatum invasion is particularly severe in dry Pittosporum undulatum Vent. (Sweet Pittosporum) is sclerophyll forest remnants where natural fire regimes a densely foliaged tree native to the wet forests of south have been altered. The suppression of natural fire re- east Australia that functions as an environmental weed gimes facilitates the invasion of some fire sensitive in habitats outside its natural range (Mullett 1996). In- invaders including P. undulatum, further compound- creasingly, some populations of P. undulatum occur- ing the displacement of fire adapted species. ring within the species natural range are also expand- Increasing cover-abundance of P. undulatum modifies ing their distribution and local densities in response to the floristic composition, structure and some functional altered ecological conditions (Rose and Fairweather elements of invaded communities and ultimately di- 1997). minishes the biodiversity values of invaded sites P. undulatum has been widely planted throughout Aus- (Mullett and Simmons 1995, Mullett 1996). P. tralia and some locations overseas for ornamental, undulatum is capable of invading a range of habitat hedge and windbreak purposes (Cooper 1956). Early types and is recognised as one of Victoria’s most seri- botanists (Maiden 1889) and gardening experts ous environmental weed species (Carr et al. 1992). (Oakman 1964) enthusiastically advocated planting of Invasion of P. undulatum into habitats outside its natu- P. undulatum for its ornamental qualities and its adapt- ral range has been listed as a ‘potentially threatening ability to a range of climatic and edaphic conditions. process’ under Schedule Three of the Victorian Flora Ironically, the same adaptive and hardy features that and Fauna Guarantee Act 1988 (Scientific Advisory so enthused early proponents of this species are now Committee 1994). regarded by conservation managers as the very 592 Twelfth Australian Weeds Conference This paper reports on some aspects of a larger study of in the germinability of P. undulatum seeds. This result P. undulatum ecology in south east Australia. also provides a relative measure of the dispersal po- tential of Pied Currawongs, which preferentially se- METHODS lect pre-dehiscent P. undulatum fruits for consump- Leaf morphology attributes were measured on fifty tion. Seeds extracted from Pied Currawong pellets and leaves sampled from 26 populations of P. undulatum Blackbird scats displayed high germinability (91.8% in south east Australia. Multivariate analyses were and 95.8% respectively), and were not significantly employed to assess the level of morphological varia- different from fresh P. undulatum seed (97.2% tion within and between populations and to investi- germinability). gate patterns of variation related to geographic and P. undulatum invasion patterns and impacts The climatic parameters. impacts of increasing cover-abundance of P, undulatum Germination trials were conducted on P. undulatum on species richness are most pronounced in the cen- seeds extracted from pre- and post-dehiscent fruits tres of radiating P. undulatum clumps. P. undulatum collected from natural and invasive populations invasion patterns and impacts were measured in sev- throughout Victoria. The germinability of P. undulatum eral natural and invaded sites however only data from seeds recovered from Blackbird scats and pellets re- one site, an invaded heathy woodland remnant on the gurgitated by Pied Currawongs were also assessed. Mornington Peninsula, Victoria, are presented. At this site, increasing cover-abundance of P. undulatum was Vegetation surveys were conducted in natural and in- significantly negatively correlated with species rich- vasive populations of P. undulatum in south east Aus- ness (r = -0.49, P <0.001) and the cover-abundance of tralia to investigate the ecological role of P. undulatum vascular plant species (r = -0.68, P <0.001). Incremen- across a range of habitats. At each site, species rich- tal increases in P. undulatum cover-abundance have a ness and the cover-abundance of vascular plants were demonstrable impact on species richness (Figure 1). recorded in 3 × 3m quadrats along 33m north-south and east-west orientated transects located through five Where P. undulatum cover-abundance levels exceed P. undulatum ‘clumps’. 20%, the competition and shading effects imposed are critical enough to exert a negative influence on some RESULTS species. Species richness declines from a mean of 15 Geographic variation in leaf morphology P. undulatum species per quadrat where P. undulatum cover-abun- leaf morphology attributes varied significantly dance is less than 20% to an average of four species in (ANOVA, all P < 0.001) across 26 populations sam- quadrats where P. undulatum cover-abundance exceeds pled in south east Australia. Leaf dimensional charac- 90%. ters (lamina length, width and distance from leaf base DISCUSSION to widest point) were significantly negatively corre- lated (all P <0.05) with altitude. Mean lamina width P. undulatum is able to adapt its phenotypic expres- was also significantly negatively correlated with lati- sion to a range of environments and environmental tude (r = -0.51, P <0.01). No clear patterns of morpho- conditions. For a species thought to have evolved in logical variation along temperature and rainfall gradi- or near wet forests it adapts extremely well to invaded ents were evident although coarse aggregations of coastal and dry forest environments. The successful populations sampled in northern New South Wales and invasion of a diverse range of vegetation complexes in South Australia were revealed through cluster analy- on other continents and islands throughout the tem- sis. The pattern of variation within and between perate, sub-tropical and tropical zones (Cooper 1956; populations sampled in Victoria was particularly com- Goodland and Healey 1996) is further evidence of the plex. species adaptability and broad ecological amplitude. Phenotypic plasticity and adaptability may compen- Dispersal and germination P. undulatum seed col- sate for low levels of genetic diversity within and be- lected from natural and invaded populations displayed tween populations of P. undulatum (Orso 1994) and high germinability. No significant differences were will facilitate further range expansion of this species recorded in the germinability of P. undulatum seeds across a range of habitats and climate regions. extracted from pre- and post-dehiscent fruits in five of the six populations sampled. This suggests that site The P. undulatum dispersal syndrome has been al- differences may be more important than fruit maturity tered in several ways since European settlement. The 593 Twelfth Australian Weeds Conference 25 between invasive fleshy-fruited spe- cies such as P. undulatum and adap- tive dispersal agents. Increased nestling predation is one 20 reported consequence of changes in the distribution, abundance and sea- sonal migration patterns of Pied 15 Currawongs in