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Ecology and consequences of invasion by non-native (wilding) in D.A. PELTZER Manaaki Whenua Landcare Research, 54 Gerald St., Lincoln 7608, New Zealand [email protected]

Abstract and many of these are considered environmental weeds. Invasion by non-native woody species into largely As a consequence, there are ongoing efforts to better treeless vegetation such as grasslands and shrublands understand and manage biological invasions at both is widespread, and has prompted both research and regional and national scales. management in response. Here I review the current Invasion by non-native species into non-forested situation of invasions by non-native Pinaceae, better areas is a global problem because tree invasions can known as ‘wilding conifers’ in New Zealand, and drive declines in conservation values or how both research and management are working to as well as altering land use suitability for agriculture better understand and manage these invaders. The or other primary industries (Richardson & Rejmánek success of wildings is explained by a combination 2011). Internationally, one of the best studied and most of history (e.g., of previously woody widespread group of invasive non-native are vegetation), biological traits of the species (rapid the pines (Pinaceae); this is unsurprising given their growth and early reproduction), and propagule pressure widespread deliberate introduction for timber, fibre (introduction effort). Wildings represent a major land and shelter (Richardson & Rejmánek 2004; Procheş use change affecting about 2 million ha, including et al. 2011). Invasive Pinaceae have become widely many grasslands, rare ecosystems and subalpine naturalised in the southern hemisphere (Richardson et habitats. Wilding invasions into grasslands have al. 1994; Simberloff et al. 2010). Most literature refers profound impacts on biological diversity, but also have to these species as invasive non-native Pinaceae or important ecosystem impacts including legacy effects conifers; but the more succinct New Zealand term is belowground by altering nutrient cycling and soil “wilding conifers” (Froude 2011). Wildings represent biota. Recent expanded efforts are underway to control about a dozen species including lodgepole and back pine and co-ordinate management to avoid or mitigate the (, P. nigra), (Pseudotsuga negative impacts of wilding conifers. The long-term menziesii) and larch (Larix decidua) (McGregor et al. value of managing invasions, and whether additional 2012). Below, I briefly review why these species are so management interventions are needed to restore successful, why they are a problem, what is currently grasslands or woody vegetation is in progress, but is being done to manage wildings, and finish with some urgently needed given recent moves to widely establish longer term or unresolved issues. new woody vegetation at large scales in New Zealand. So why are wildings so successful? Keywords: biological invasions, ecological impacts, Successful establishment, naturalisation and spread of ecosystem legacies, non-native tree species, Pinus introduced species is often attributed to their biological contorta, plant-soil interactions, removal effects, characteristics such as rapid growth, high reproductive wilding conifers output and tolerance of environmental conditions. For example, lodgepole pine (Pinus contorta) grows Introduction much faster in countries where it has been introduced Biological invasions by non-native species are an compared to the home range in North America important component of global change, and are a (Ledgard 2001; Taylor et al. 2016a). However, for pine major driver of changes in ecological communities and species introduced to New Zealand the best predictors ecosystems (Vitousek et al. 1997; Early et al. 2017). of naturalisation are introduction effort ( use, Moreover, many of these drivers interact, for example, area planted, and date of introduction) and climate- changing land use or climate can exacerbate the number matching; the main biological trait is early age to or success of biological invaders in a system (Tylianakis reproduction (McGregor et al. 2012). In addition, et al. 2008). More specifically, New Zealand is one of seed traits (terminal velocity, or the rate of seed fall the world’s most invaded countries. This is highlighted vertically) and wind, interact to drive both short and through such national-scale efforts as predator-free long-distance dispersal of conifers, and in combination New Zealand. For plants, about half of the New with high survivorship of seedlings, this allows pine Zealand flora is comprised of non-native plant species, invasions to rapidly establish in grasslands (Buckley et

ISSN 2463-2872 (Print) ISSN 2463-2880 (Online) 40 Journal of New Zealand Grasslands 80: 39-46 (2018) al. 2005; Caplat et al. 2012). Overall, this demonstrates that early, large-scale plantings overcome the lag times between when a species is introduced and when it becomes naturalised and invasive, but also suggests that some species that are not currently widespread invaders are able to spread, but are just taking longer to invade because of lower initial introduction effort. Positive interactions can also occur among introduced species. For conifers, the most obvious is mutualisms involving mycorrhizas: beneficial fungi associated with plant roots that are needed for tree establishment and can improve the subsequent survival and growth of plants. For example, lodgepole pine co-invades with non-native mycorrhizal fungi rather than associating with native fungi (Dickie et al. 2010), and even a single mycorrhizal species can enable pine invasion (Hayward et al. 2015). Furthermore, few mycorrhizal fungal species are needed to facilitate pine invasion, including into grasslands that do not contain ectomycorrhizal tree species. This differs from native beech (nothofagus) tree species that are also ectomycorrhizal, but are notoriously slow at reinvading grasslands. In addition, non-native mycorrhizal fungi can be transported by non-native mammals such as deer and possums, but presumably other mammals can also disperse fungi ( et al. 2015). Positive interactions amongst wilding tree species, their co- invading mycorrhizal fungi and dispersal of these fungi by non-native mammals helps to explain the invasion success of wilding conifers into grasslands (Figure 1).

Why are wildlings a problem? The invasion by trees into grassland has both obvious consequences and less obvious legacies. Although the invasion potential of conifers has been known for some time (e.g., Benecke 1967; Hunter & Douglas 1984; and references in Froude 2011), wildings are now broadly viewed as a serious threat to grassland biodiversity, pasture or primary production, and some ecosystem processes, through profoundly altering vegetation composition and resource availability (Dickie et al. Figure 1 Invasion by non-native pines or ‘wilding conifers’. Early invasion in the Mackenzie Basin (top panel), 2011; Rundel et al. 2014; Taylor et al. 2016b). Different ongoing invasion near Hanmer Springs (middle impacts of wildings occur at differentFigure stages 1 Invasion of the by non-nativepanel), pines and or ‘wildinga long-invaded conifers’. site in theEarly Kawekas invasion in the invasion process. Grassland plant diversityMackenzie can initially basin (top panel), ongoing(bottom invasionpanel). near Hanmer Springs (middle panel), and a increase with invasion but then rapidly declines; carbon long-invaded site in the Kawekas (bottom panel). scales positively and linearly with increasing tree following the control of wildings (Dickie et al. 2014). density; and water yield declines most rapidly with This includes increases in both soil available nitrogen canopy closure because of rainfall interception (Fahey and phosphorus compared to native grasslands, and

& Payne 2017). As a consequence, different impacts of changes in plant-associated nematodes that can act as wildings occur as invasion proceeds, and this can create pathogens. What is surprising is how rapidly impacts trade-offs in net impacts, or conversely, the benefits and legacies can develop (<10 years) given the invasion of managing invasions for conservation or primary is by tree species (Figure 1; Taylor et al. 2016a). 13 production (Mason et al. 2017). The less obvious The total area affected by wildings (defined as a legacies of wildings include belowground changes minimum of one tree/ha) is thought to be around 2 to nutrient cycling and the soil biota that can persist million ha, or about the same area as is in Ecology and consequences of invasion by non-native (wilding) conifers in New Zealand (D.A. Peltzer) 41 (MPI 2014). The area invaded has by some estimates increased by about 5%/year over the past few decades. Because wildings can invade high-country grasslands, threaten rare ecosystems (many of which are treeless vegetation and have high conservation value), and can establish above the natural treeline, they are considered a national-scale weed issue (Froude 2011). The speed and scale of wilding invasion occurred despite ongoing management efforts by landowners, community trusts and government departments alike, and this prompted a call for increased management to contain or eradicate wildings. In response, the Ministry of Primary Industry spearheaded FigFigureure 2 Illustrative 2 Illustrative state and transition state model and for grasslandstransition modified model and simplifiedfor from a national strategy to raise general awareness of this Standish et al. (2009).grasslands Arrows show modified vegetation andchanges simplified from tussock from grassland Standish towards issue, and to co-ordinate management efforts amongst dominance by eitheret native al. (2009).(left side) orArrows non-native show weeds vegetation (right side). Dependingchanges on propagule pressure from(seed sources)tussock and grassland environment, towardsmost trajectories dominance tend towards by central and regional government, landowners and dominance by non-native trees or wilding conifers, but in some cases states can persist either native (left side) or non-native weeds (right industry at a national scale (MPI 2014). More recently, through self-replacement (shown by loops for a vegetation-type). side). Depending on propagule pressure (seed this has prompted formation of the National Wilding sources) and environment, most trajectories tend Management Programme that organises and towards dominance by non-native trees or wilding co-ordinates these renewed management efforts (www. conifers, but in some cases states can persist wildingconifers.org.nz). through self-replacement (shown by loops for a removal vegetation-type). c) ecosystem post-tipping point What is currently being done to manage wildings? strategies have contrasting outcomes for vegetation

In response to widespread invasion, there are many recovery. Forn example, early control of wilding io vas management programs internationally that are pursued seedling orin saplings returned a tussock grassland b) secondarily invaded ecosystem to prevent or mitigate the spread and impacts of community to resemble the uninvaded community, additional biological invaders (e.g., Hulme 2015; Nuñez et al. whereas later removal of trees droveinterventions the vegetation

uninvad ed 2017). Usually, multiple management strategies are towardsec osystem dominance by few non-native grasses (e.g., deployed in an attempt to reduce the abundance or Agrostis capillaris, Anthoxanthum odoratuma); restoredLedgard ec osystem distribution of , but with an ultimate & Paul 2008; Paul & Ledgard 2009; Dickie et al. goal of avoiding or mitigating negative impacts on 2014). Overall, woody invasion into grasslands follows native ecosystems (e.g., Zavaleta et al. 2001; Barney somewhat predictable state and transitions (Figure et al. 2015). For wilding conifers there are multiple 2), but the rapid shift towards dominance of wilding 14 methods used for controlling trees that are used conifers rather than native woody plant species is a depending on tree density, the species involved and major concern (Standish et al. 2009; Young et al. 2016). site access. These include mechanical (grubbing, More specifically, management of wildings can have pulling, cutting, mulching) and chemical (ground and different outcomes depending on the methods used aerial basal bark herbicides, aerial boom spraying) and the more rapid establishment of many non-native methods as well as land management (mob stocking, plants compared to native species (Wardle & Peltzer pasture improvement). The selection and use of 2017). Other potential management tools including different management strategies is well-covered in fire, biological control and the use of sterile trees in work plans, user guides and vast previous experience , are all options for efficient longer-term (e.g., Ledgard 2009; Briden et al. 2014; Gous et al. management to prevent future wilding spread, but are 2014, 2015). Current total spending nationally wilding not currently in use. control is about $16 million/year, but this excludes Many invasive plant management or eradication ongoing management efforts on private land and by efforts in general either fail to meet their stated communities (MPI 2014). management goals or do not carry out rigorous Selection of different management strategies typically evaluations of success (e.g., Maxwell et al. 2009; considers the effectiveness of control (e.g., proportion Wilson et al. 2014); those that do measure effectiveness of trees killed) and cost, and in some cases, the response as change in invasive species to management per se of native communities (Flory & Clay 2009). A few rather than wider values or benefits (Reid et al. 2009). studies that consider the non-target impacts of wildings However, an implicit goal of management, including show that changes in plant communities caused by both of wildings, is often to generate benefits beyond wilding invasion and different management or removal immediate control of the invader itself. These benefits 42 Journal of New Zealand Grasslands 80: 39-46 (2018)

Figure 3 General trajectories of changes driven by invasion, invader removal or restoration. Management (‘removal’) of invaders can involve creating a new system (top right), reinvasion by wildings or secondary invasion by other weeds (middle right), or with sufficient seed sources or additional management, restoration towards an uninvaded system (bottom left) (modified from Wardle & Peltzer 2017). span conservation of sites and native species, sustained 2. If eradication or elimination of wilding conifers is not water yield in catchments, and maintaining landscape possible for a site, then what is the minimal wilding values (MPI 2014; Nuñez et al. 2017). abundance that management should achieve to ensure benefits within a site (Barney et al. 2015)? In other Looking ahead words, can sustaining a low abundance of wildings Non-native conifers in New Zealand are an interesting provide nearly all of the benefits of management? issue globally because they both underpin the plantation 3. Taking a longer-term view, are additional management industry, but at the same time most species are interventions needed to establish grassland or native also considered environmental weeds. The two issues woody species (e.g., McAlpine et al. 2016), or to are not directly comparable, however, because both the prevent undesirable ‘secondary invasions’ by other tree species involved and locations can differ between non-native species (Figure 3; see also Firn et al. plantation forestry and wilding invasion. Overall, 2010)? The long-term options for wilding invaded wildings are currently a widespread weed problem of sites converge on active management towards non-native trees invading largely into grasslands and grassland (either native tussock or pasture), or woody other non-forest vegetation-types, but new progress in vegetation (most likely mixtures of native and non- management to contain or eradicate wildings is now native woody species); making the end-point explicit being made through greater co-ordination and operational is necessary to know when and what additional funding at a national scale (http://wildingconifers.org.nz/ management efforts are needed. about-us/programme-2/). However, there remain several 4. What is the future risk of wilding invasion given opportunities or gaps in understanding that remain to be changes in land use, climate and management? This filled, and that closely linking management and research is particularly pressing given that ongoing, large- can help to resolve including: scale changes in land use are likely (e.g., driven 1. There is no one-off management method that can by the billion trees Government initiative), and the eliminate wildings in perpetuity, but rather follow- prediction that future climates might alter both plant up or repeated management is typically needed performance (growth, height and reproductive effort) (Ledgard 2009; Nuñez et al. 2017). Determining how and extreme events (e.g., storms), thus potentially long management must endure to control or contain increasing spread risk (Caplat et al. 2013). wildlings is needed, but planning for a decade or Controlling invasive non-native species such as wilding longer is often essential. conifers is at the same time both simple (i.e., killing Ecology and consequences of invasion by non-native (wilding) conifers in New Zealand (D.A. Peltzer) 43 unwanted trees) and complex because it involves Dickie, I.A.; Yeates, G.W.; John, M.G.S.; Stevenson, manipulation of complex systems across all stages of B.A.; Scott, J.T., Rillig, M.C.; Peltzer, D.A.; Orwin, the invasion process from preventing new invasions, to K.H.; Kirschbaum, M.U.; Hunt, J.E.; Burrows, L.E. managing established populations, and in a few cases, 2011. and biodiversity trade- sorting out long-invaded areas. Because the outcomes offs in two woody successions. Journal of Applied of invasive species removal, in terms of conservation Ecology 48: 926-934. values and ecosystem impacts, are often variable Dickie, I.A.; St John, M.G.; Yeates, G.W.; Morse, (Kettenring & Adams 2011), it is essential to evaluate C.W.; Bonner, K.I.; Orwin, K.; Peltzer, D.A. 2014. the performance of different management strategies. 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