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Naturalised Plants Transform the Composition and Function of the New Zealand flora

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Naturalised Plants Transform the Composition and Function of the New Zealand flora Biol Invasions (2021) 23:351–366 https://doi.org/10.1007/s10530-020-02393-4 (0123456789().,-volV)( 0123456789().,-volV) ALIEN FLORAS AND FAUNAS 7 Naturalised plants transform the composition and function of the New Zealand flora A.J. Brandt . P.J. Bellingham . R.P. Duncan . T.R. Etherington . J.D. Fridley . C.J. Howell . P.E. Hulme . I. Jo . M.S. McGlone . S.J. Richardson . J.J. Sullivan . P.A. Williams . D.A. Peltzer Received: 16 March 2020 / Accepted: 15 October 2020 / Published online: 29 October 2020 Ó Springer Nature Switzerland AG 2020 Abstract The New Zealand flora has a high propor- strategy. The 1798 species in the naturalised flora tion of endemic species but has been invaded by currently comprise 43.9% of the total number of almost the same number of non-native plant species. vascular plant species, and add 67 plant families and To support management of invasive plant species, we 649 genera to the total vascular flora. The naturalised provide an updated inventory of New Zealand’s flora has a greater proportion of herbaceous species naturalised flora and compare it with the native flora and annual species than the native flora, which could to identify key taxonomic and functional distinctions. influence ecosystem processes such as decomposition We also assess how the naturalised flora may impact and nutrient cycling. Naturalised trees have higher leaf ecosystem processes differently than the native flora nitrogen concentration for a given leaf area than native using functional traits related to plant resource use trees, which could increase rates of nutrient cycling in invaded forest ecosystems. A greater number of naturalised species are present in larger, more north- Electronic supplementary material The online version of this article (https://doi.org/10.1007/s10530-020-02393-4) con- erly, and more populated regions of New Zealand. Our tains supplementary material, which is available to authorized results demonstrate both taxonomic and functional users. A.J. Brandt (&) J.D. Fridley Manaaki Whenua – Landcare Research, Dunedin, New Department of Biology, Syracuse University, Syracuse, Zealand NY, USA e-mail: [email protected] C.J. Howell P.J. Bellingham Á T.R. Etherington Á Department of Conservation, Wellington, New Zealand I. Jo Á M.S. McGlone Á S.J. Richardson Á P.A. Williams Á D.A. Peltzer P.E. Hulme Manaaki Whenua – Landcare Research, Lincoln, New Bio-Protection Research Centre, Lincoln University, Zealand Lincoln, New Zealand P.J. Bellingham J.J. Sullivan School of Biological Sciences, University of Auckland, Department of Pest Management and Conservation, Auckland, New Zealand Lincoln University, Lincoln, New Zealand R.P. Duncan Institute for Applied Ecology, University of Canberra, Bruce, ACT, Australia 123 352 A.J. Brandt et al. differences between the native and naturalised flora of immigration (McGlone et al. 2001), and the extant New Zealand that can be used to guide management of flora is surprisingly young, mostly arriving after the naturalised plants, including the 314 species currently Oligocene (Heenan and McGlone 2019). The flora is managed as environmental weeds, from the local to also functionally distinct compared to other temperate, national scale. continental areas, a consequence of its isolation and highly oceanic climatic regime (McGlone 2006). Prior Keywords Exotic species Á Functional traits Á to human settlement, dense evergreen podocarp- Indigenous plant species Á Naturalised plant species Á angiosperm forests dominated lowland to subalpine Plant growth forms Á Regional distribution landscapes. Few native species are deciduous or summer-green (McGlone et al. 2004) and many native tree species are small-statured (McGlone et al. 2010). Mammalian browsers and grazers were completely Introduction absent prior to European arrival (Atkinson 2006), their place taken by birds which, lacking teeth and prehen- New Zealand is an isolated archipelago ranked among sile tongues, may exert quite different pressures on the the world’s top 25 biodiversity hotspots (Myers et al. vegetation (Antonelli et al. 2010; Wilson and Lee 2000) because of high endemism (e.g., 83% of 2012). These and other differences have resulted in a reptiles, 99% of millipedes, 100% of conifers, 85% flora lacking species with trait combinations that are of angiosperms; Lee, Lee 2015) but it is also ‘‘ravaged common in other similar temperate environments; by biological invasions’’ (Simberloff 2009). The such apparently unoccupied ecological spaces have effects of invasive non-native mammals in New been described as ‘functional gaps’ (Blonder 2016; Zealand are well documented, and introduced mam- Dansereau 1964; Lee 1998). mals are the focus of national initiatives to reduce or The introduction of non-native plant species to New even eradicate some species (Owens 2017; Peltzer Zealand by humans broke down biogeographic barri- et al. 2019). However, a contemporary nationwide ers and filled apparent functional gaps in the flora. management strategy for non-native plants has lagged When Ma¯ori arrived in New Zealand (ca. 1280 AD; behind that developed for mammals, despite legisla- Wilmshurst et al. 2008), approximately 90% of the tive efforts (reviewed in Hulme 2020) to control the archipelago was forested (McGlone 1983) and open high levels of plant invasions (Gatehouse 2008; habitats were confined to the alpine and other areas Williams and West 2000) that continue to increase where disturbance, climate or soils precluded domi- through ongoing naturalisation (i.e. establishment of nance by trees. Fire frequency and severity increased self-sustaining wild populations of introduced non- significantly with human settlement, transforming native species, Blackburn et al. 2011; Howell 2019; large areas of closed canopy forest ecosystems to Williams and Cameron 2006). Here, we update a open, seral vegetation (McGlone 1983; McWethy previous inventory of the naturalised flora of New et al. 2010). Colonisation by Europeans from the Zealand (Howell and Sawyer 2006) and compare it 1770s onwards accelerated deforestation. Fire regimes with the native flora to identify key taxonomic and were intensified, wetlands were drained, and most functional distinctions. Our aim is to support man- suitable soils were given over to pastoral agriculture. agement of naturalised plants at local and national Both pastoralism and forestry were initially based on scales by summarising our current knowledge of native grasses and trees but quickly transitioned to a naturalised species traits and distribution at the scale near complete dependence on non-native species. of the whole flora (Blackburn et al. 2014; Nunez-Mir More than half of New Zealand’s land area is now et al. 2019). comprised of a range of ecosystems dominated by The native flora is highly endemic at the species non-native plants (Hulme 2020; Thomson 1922; level but very few genera and no plant families are Walker and Bellingham 2011). endemic to New Zealand. The ca. 2000 km of ocean Non-native plant species were introduced to New that has separated New Zealand from the nearest Zealand deliberately for agriculture, forestry and continent (Australia) for at least 50 million years has horticulture (Gatehouse 2008; Smith et al. 2000; acted more as a selective filter than a barrier to Thomson 1922; Williams and Cameron 2006), with 123 Naturalised plants transform the composition and function of the New Zealand flora 353 accidental introductions accompanying many of these could result in naturalised species having substantial sources (e.g. as seed contaminants). Deliberately environmental and economic impacts. For example, introduced species were selected for their ornamental naturalised grasses and herbs are often faster-growing novelty or to fill functions or services not provided and shorter-lived, with lower tissue density and higher sufficiently by native species (see Castro-Dı´ez et al. foliar nitrogen (N) concentrations than native species 2019 for a global treatment of this trend; Hulme 2020). (Craine and Lee 2003; Kichenin et al. 2013; Peltzer Ma¯ori introduced perhaps a dozen plant species from et al. 2009), with consequences for grassland commu- the Pacific (Horrocks 2004; Leach 2005; Williams and nity dynamics (Gross et al. 2013). Additionally, Cameron 2006), while Europeans introduced[25,000 several naturalised woody species have traits linked species from all regions of the world (Crosby 2004; to pyrogenicity such as serotiny, thick bark, the ability Diez et al. 2008, 2009; Fridley and Sax 2014; to sprout vigorously, and high litter (fuel) accumula- Gatehouse 2008; Howell 2019). Deliberately intro- tion (Wyse et al. 2018), traits which few species in the duced species were often selected for their ability to native flora possess because natural fires were rare in thrive under the prevailing climatic conditions, ease of New Zealand prior to human colonisation (Perry et al. establishment, rapid growth, and high fecundity, and 2015). Greater understanding of functional differences unintentionally introduced species include cosmopoli- between the native and naturalised flora should better tan invaders with rapid growth and high fecundity enable land managers to prioritise management efforts (e.g., Jacobaea vulgaris, Erigeron sumatrensis). It is based on likely impacts (e.g. faster nutrient cycling thus unsurprising that ca. 10% of plant species and litter decomposition associated with higher foliar introduced to New Zealand have naturalised (i.e. N). established wild populations; Williams and Cameron Here,
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