Compared to natural forests, fast-growing plantations of exotic species such as Pinus radiata are often perceived as marginal habitat or unsuitable habitat for most native species. By studying Coleoptera (beetles) in a variety of landscape elements (pasture, native forest and different aged Pinus radiata stands) in a highly modified and fragmented landscape in New Zealand I aimed to determine the value of exotic plantation forests for native biodiversity, and how these species are affected by different sized clearfell harvest areas. Pitfall trap sampling of beetles showed that plantation forest stands can provide suitable complimentary habitat to native forest for many species. Rarefied species richness of Carabidae, Scarabaeidae and Scolytinae was not significantly different between habitats, however, habitat types differed significantly in their beetle community composition. Comparing different production habitats, Pinus radiata stands had a beetle community composition most similar to native forest. However, a small minority of species, e.g., Dichrochile maura, were restricted to native forest habitat highlighting the importance of retaining indigenous ecosystems within plantations. Unlike human modified habitats, native forests did not provide suitable habitat for exotic species. Clearfell harvesting is controversial and its impact on biodiversity is a key constraint for many forest certification programs, such as that administered by the Forest Stewardship Council (FSC). Despite this, no replicated manipulative experimental studies of the impact of different sized clearfell harvest areas on biodiversity have been undertaken at scales relevant to the New Zealand forest industry. One potential model of the impact of different clearfell harvest sizes is the concept of a threshold size. A threshold scenario may occur where clearfell harvest impacts increase at a rate disproportionate to the change in clearfell size over a small range of harvest areas, but impacts remain relatively unchanged either side of the threshold zone. I sampled Coleoptera in experimentally created 0.01, 0.05, 0.5, 5.0, 50 and 500 ha clearfells within Pinus radiata plantations in the central North Island of New Zealand. The wide range of clearfell harvest sizes, including some very small areas, such as 0.01 ha was instigated in an attempt to document potential clearfell harvest size thresholds. Rarefied native beetle species richness was higher in harvest areas compared to adjacent mature plantation stands. The beetle species richness in 5 ha and 500 ha harvest areas was significantly greater species than that in small 0.01 - 0.5 ha harvest areas. Although, the high beetle diversity recorded in 500 ha clearfells should be treated with caution due to confounding spatial autocorrelation. The degree of change in beetle community composition increased with increasing clearfell harvest area. Beetle assemblages in large harvest areas were less similar to their paired adjacent mature forest than smaller harvest areas. Although, constrained multivariate ordination techniques did show a short-term change in beetle species composition between recently clearfelled harvest areas of as little as 0.05 ha and adjacent mature P. radiata stands. The colonisation by open-habitat disturbance-adapted species was a key driver of this change, some species dispersed into clearfelled stands in significant densities within days post-harvest. Overall, there were no distinct short-term trends to the change in species richness as a function of increasing harvest area that would suggest an ecological impact threshold response. If short-term outcomes of clearfell harvesting are ameliorated by successful recolonisation, the long-term spatial arrangement of different aged stands becomes more important for the maintenance of biodiversity at the landscape level than short-term consequences of harvesting. By sampling selected beetle taxa in 1, 2, 4, 8, 16 and 26 year-old stands, I found that the abundance of seven out of eight of the species selected for analysis recovered to levels similar to those in adjacent mature forest within the timeframe of a single harvest rotation. Individual species utilised different aged stands, indicating different life-history strategies. For example, open-habitat, disturbance-adapted species such as Cicindela tuberculata and Sitona discoideus were prominent in young stands, and forest species such as Pycnomerus sophorae and Paracatops phyllobius were highly abundant in older stands. These alternative life-history strategies highlight the benefits of maintaining a mixture of different aged stands to increase biodiversity at the landscape level. This thesis fills an important gap in our knowledge of biodiversity in production landscapes. I show that plantation forests have value as complimentary habitat to native forest and they make an important contribution to the maintenance of biodiversity at the landscape level. Although clearfell harvesting is a severe disturbance to the forest ecosystem, the long-term recovery of beetle populations suggests that harvesting is not the key limiting factor to the enhancement of biodiversity in the plantation forests studied. This unusual situation is possibly the result of prior land-use history, as many plantations were established on degraded pastoral land, and harvest-sensitive species are unlikely to have survived this initial land-use change. As such, the severity of the long-term impacts of clearfell harvesting on biodiversity are likely to be context specific and will vary accordingly. The importance of spatial heterogeneity of habitat elements, including different aged plantation stands and native forest remnants, needs to be investigated in more detail to determine what limits biodiversity in this plantation landscape. Key points to consider are the proximity to, and proportion of, native forest cover in the landscape and the degree of connectivity among native remnants. It is these landscape-level attributes that may determine biodiversity at a regional scale, and more emphasis should be placed on landscape scale factors and there interaction with stand specific forest management practices. For example, the spatial mosaic of harvesting areas may need to be of a finer-scale when there are fewer native remnants within the landscape.

Compared to natural forests, fast-growing plantations of exotic species such as Pinus radiata are often perceived as marginal habitat or unsuitable habitat for most native species. By studying Coleoptera (beetles) in a variety of landscape elements (pasture, native forest and different aged Pinus radiata stands) in a highly modified and fragmented landscape in New Zealand I aimed to determine the value of exotic plantation forests for native biodiversity, and how these species are affected by different sized clearfell harvest areas. Pitfall trap sampling of beetles showed that plantation forest stands can provide suitable complimentary habitat to native forest for many species. Rarefied species richness of Carabidae, Scarabaeidae and Scolytinae was not significantly different between habitats, however, habitat types differed significantly in their beetle community composition. Comparing different production habitats, Pinus radiata stands had a beetle community composition most similar to native forest. However, a small minority of species, e.g., Dichrochile maura, were restricted to native forest habitat highlighting the importance of retaining indigenous ecosystems within plantations. Unlike human modified habitats, native forests did not provide suitable habitat for exotic species. Clearfell harvesting is controversial and its impact on biodiversity is a key constraint for many forest certification programs, such as that administered by the Forest Stewardship Council (FSC). Despite this, no replicated manipulative experimental studies of the impact of different sized clearfell harvest areas on biodiversity have been undertaken at scales relevant to the New Zealand forest industry. One potential model of the impact of different clearfell harvest sizes is the concept of a threshold size. A threshold scenario may occur where clearfell harvest impacts increase at a rate disproportionate to the change in clearfell size over a small range of harvest areas, but impacts remain relatively unchanged either side of the threshold zone. I sampled Coleoptera in experimentally created 0.01, 0.05, 0.5, 5.0, 50 and 500 ha clearfells within Pinus radiata plantations in the central North Island of New Zealand. The wide range of clearfell harvest sizes, including some very small areas, such as 0.01 ha was instigated in an attempt to document potential clearfell harvest size thresholds. Rarefied native beetle species richness was higher in harvest areas compared to adjacent mature plantation stands. The beetle species richness in 5 ha and 500 ha harvest areas was significantly greater species than that in small 0.01 - 0.5 ha harvest areas. Although, the high beetle diversity recorded in 500 ha clearfells should be treated with caution due to confounding spatial autocorrelation. The degree of change in beetle community composition increased with increasing clearfell harvest area. Beetle assemblages in large harvest areas were less similar to their paired adjacent mature forest than smaller harvest areas. Although, constrained multivariate ordination techniques did show a short-term change in beetle species composition between recently clearfelled harvest areas of as little as 0.05 ha and adjacent mature P. radiata stands. The colonisation by open-habitat disturbance-adapted species was a key driver of this change, some species dispersed into clearfelled stands in significant densities within days post-harvest. Overall, there were no distinct short-term trends to the change in species richness as a function of increasing harvest area that would suggest an ecological impact threshold response. If short-term outcomes of clearfell harvesting are ameliorated by successful recolonisation, the long-term spatial arrangement of different aged stands becomes more important for the maintenance of biodiversity at the landscape level than short-term consequences of harvesting. By sampling selected beetle taxa in 1, 2, 4, 8, 16 and 26 year-old stands, I found that the abundance of seven out of eight of the species selected for analysis recovered to levels similar to those in adjacent mature forest within the timeframe of a single harvest rotation. Individual species utilised different aged stands, indicating different life-history strategies. For example, open-habitat, disturbance-adapted species such as Cicindela tuberculata and Sitona discoideus were prominent in young stands, and forest species such as Pycnomerus sophorae and Paracatops phyllobius were highly abundant in older stands. These alternative life-history strategies highlight the benefits of maintaining a mixture of different aged stands to increase biodiversity at the landscape level. This thesis fills an important gap in our knowledge of biodiversity in production landscapes. I show that plantation forests have value as complimentary habitat to native forest and they make an important contribution to the maintenance of biodiversity at the landscape level. Although clearfell harvesting is a severe disturbance to the forest ecosystem, the long-term recovery of beetle populations suggests that harvesting is not the key limiting factor to the enhancement of biodiversity in the plantation forests studied. This unusual situation is possibly the result of prior land-use history, as many plantations were established on degraded pastoral land, and harvest-sensitive species are unlikely to have survived this initial land-use change. As such, the severity of the long-term impacts of clearfell harvesting on biodiversity are likely to be context specific and will vary accordingly. The importance of spatial heterogeneity of habitat elements, including different aged plantation stands and native forest remnants, needs to be investigated in more detail to determine what limits biodiversity in this plantation landscape. Key points to consider are the proximity to, and proportion of, native forest cover in the landscape and the degree of connectivity among native remnants. It is these landscape-level attributes that may determine biodiversity at a regional scale, and more emphasis should be placed on landscape scale factors and there interaction with stand specific forest management practices. For example, the spatial mosaic of harvesting areas may need to be of a finer-scale when there are fewer native remnants within the landscape.

Plantations are widespread throughout temperate regions, and the area of plantation land cover is predicted to get larger in the future. Interest in ensuring sustainable plantation management is also growing, as it is increasingly recognised that productive areas should play a role in biodiversity conservation. Plantation landscapes can comprise the majority of forested cover in some countries, but taxon-specific guidance can be lacking, due to plantations often being under surveyed. Therefore, despite substantial incentives existing to ensure that plantations meet various ecological criteria, plantation managers lack the information necessary to implement effective management plans. Many bat species have undergone widespread declines in recent decades, attributed to habitat loss and fragmentation, particularly of forested habitat. In many temperate countries, historical deforestation has resulted in very low native tree cover, and subsequently, considerable replanting with non-native commercial coniferous plantations has taken place. Species specific habitat surveys have often demonstrated avoidance of conifer plantations by bats, which has been attributed to a lack of roosts and low invertebrate prey abundance. Furthermore, widespread lepidopteran declines have been partly attributed to afforestation with non-native conifer, but moth associations with commercial coniferous plantations are usually only studied for pest species. Bats present a particular challenge in plantation landscapes; tree cover is important to many species to a greater or lesser extent, and in the United Kingdom, destruction of a roost site is illegal, regardless of whether it was deliberate or accidental. However, the extent to which bats associate with non-native commercial plantations is relatively unexplored. This is the first study to explicitly test bat associations with Picea sitchensis plantations (using acoustic detectors, trapping and radio tracking), and shows that, contrary to expectations, they may be an important habitat for breeding populations of Pipistrellus spp., particularly P. pygmaeus. High levels of activity were recorded for both P. pygmaeus and P. pipistrellus, despite little difference in dipteran abundance between different stand types, both species preferentially foraged in felled or less dense stands. This suggests that bats preferentially forage in areas with less acoustic and physical clutter, which will increase foraging efficiency. The impacts of felling in non-native commercial coniferous plantations on foraging activity was tested, for the first time, using a Before – After – Control – Impact experimental design. Bat activity (specifically P. pipistrellus and Nyctalus) increased after felling, particularly in smaller stands. In contrast felling had significant, negative impacts on moth abundance, species richness and diversity, and these effects remain after constraining for functional trait similarity. Reductions in richness and diversity in response to felling were similarly large for both rare and abundant species. Therefore, while bats may benefit from clear fell practices, albeit as long as the size of patches is small, moth populations could benefit from a shift towards other forestry methods, iv such as continuous cover forestry. These results also have implications for the recent, but increasing practice of siting wind turbines in commercial coniferous plantations, as pre-installation preparation involves clearing small patches of forest which may attract foraging bats; post felling monitoring should be carried out to examine potential impacts on bat populations. The presence of broadleaf trees in and around plantations significantly increased moth richness, mostly through increased occurrence of rare species. Broadleaf woodlands (defined as land spanning more than 0.5 ha, with trees higher than 5m and a combined cover of shrubs, bushes and trees above 10%), also had higher functional redundancy than plantation sites. For a diverse moth population to persist in plantation landscapes, preserving remnant patches of broadleaf trees is essential. There was little difference in bat activity between broadleaf woodlands and plantation sites. However, bat abundance, particularly that of reproductively active females, was greater in broadleaf sites compared to plantations. This was particularly true for Myotis and Nyctalus spp., very few of which were trapped in commercial plantations. Therefore, although reproductively active female Myotis bats are present in the surrounding landscape, they do not appear to associate with plantations themselves. This may reflect a lack of roost availability; both P. pygmaeus and P. pipistrellus preferentially form large maternity colonies in buildings, but for Myotis and Nyctalus spp. which roost switch regularly and often use trees, it is unlikely many suitable roosts exist within the plantations themselves. Many substantial P. pygmaeus maternity colonies were identified in and around Galloway forest, with some holding more than 500 individuals. All maternity colonies were in buildings, and most inhabited (and one uninhabited) buildings within the plantation contained a roost. Although females occasionally used old or dead deciduous trees as temporary roosts, there was no evidence of roosting in crop trees such as P. sitchensis. During this study, the Forestry Commission installed 36 bat boxes; within 6 months over 90% had been used, with a number of harems found inside. This fast uptake compared with bat use of boxes in other locations reflects the paucity of appropriate structures for either roost or harem use in commercial plantations. Twelve bats were captured while foraging, tagged with small radio transmitters, and followed for between 2 and 6 nights during 2014 and 2015. All but one tagged female preferentially foraged within the plantation, with individuals selecting equally riparian habitats and felled stands. Tagged females which roosted furthest from the plantation had the largest home ranges; one individual flew nearly 40km each night to reach foraging areas distant from her roost, suggesting that the food availability within the plantation was sufficient to render such a long journey energetically viable. v These results have important implications for bat populations in and around commercial coniferous plantations. Far from being avoided by bats, plantation landscapes may constitute an important habitat type for both P. pygmaeus and P. pipistrellus, likely due to the high abundance of nematoceran diptera in plantation woodlands. Furthermore, plantation forests support a similar richness of moth species to urban and agricultural woodlands, including a number of declining species of special conservation concern. A list of management recommendations to benefit both bat and moth populations in commercial plantations is presented at the end of this thesis.