Evolution of New Zealand Insects: Summary and Prospectus for Future Re- Search

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Buckley, Thomas, Krosch, Matthew, & Leschen, Richard (2015) Evolution of New Zealand insects: summary and prospectus for future research. Austral Entomology, 54(1), pp. 1-27.

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The evo lution of New Zealand insects: summary and prospectus for future research

Journal:For Austral Review Entomology Only Manuscript ID: Draft

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Complete List of Authors: Buckley, Thomas; Landcare Research, Krosch, Mathew; The University of Queensland, Centre for Water in the Minerals Industry Leschen, Richard; Landcare Research,

Keywords: Biogeography, adaptation, alpine, freshwater, island

Knowledge on the evolution of the New Zealand insect fauna is reviewed and outstanding questions are highlighted. The New Zealand insect fauna is a composite of old and recent lineages and many spectacular examples of evolutionary processes are evident including; adaptive radiation, hybridisation, and unusual adaptations. We discuss the origins and evolution of four key communities within the insect fauna: terrestrial lowland insects, alpine insects, aquatic insects, and insect communities from offshore islands. Within each of these communities key lineages are discussed and in particular the key adaptations that enable these species Abstract: to thrive. Glacial history has had a dramatic impact on the New Zealand insects, and the effects on different lineages are discussed. The New Zealand insects are unique, yet many are threatened with extinction and efforts to preserve the fauna are reviewed. Despite the accumulating knowledge, major gaps still exist and these are outlined, as are opportunities to address key questions. The review ends with a synthesis and a discussion of how systematics, new technologies, and integrative approaches have the promise to improve dramatically our understanding of New Zealand insect evolution.

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1 1 2 3 4 The evolution of New Zealand insects: summary and 5 6 7 prospectus for future research 8 9 10 11 12 13 14 Thomas R. Buckley 1,2,3,* 15 16 4 17 Matt Krosch 18 For Review Only 19 Richard A.B. Leschen 1 20 21 22 23 24 1 25 Landcare Research, Private Bag 92170, Auckland, New Zealand 26 27 2School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland, New 28 29 Zealand 30 31 3 32 Allan Wilson Centre, Auckland, New Zealand 33 34 4 Centre for Water in the Minerals Industry, Sustainable Minerals Institute, The University of 35 36 Queensland, Brisbane, Queensland 4072, Australia 37 38 39 40 41 *[email protected] 42 43 44 45 46 47 Running title: Evolution of New Zealand insects 48 49 50 51 52 53 54 55 56 57 58 59 60 Austral Entomology Page 2 of 81

2 1 2 3 Abstract 4 5 6 Knowledge on the evolution of the New Zealand insect fauna is reviewed and outstanding 7 8 questions are highlighted. The New Zealand insect fauna is a composite of old and recent 9 10 lineages and many spectacular examples of evolutionary processes are evident including; 11 adaptive radiation, hybridisation, and unusual adaptations. We discuss the origins and evolution 12 13 of four key communities within the insect fauna: terrestrial lowland insects, alpine insects, 14 15 aquatic insects, and insect communities from offshore islands. Within each of these communities 16 17 key lineages are discussed and in particular the key adaptations that enable these species to 18 thrive. Glacial history Forhas had a dramaticReview impact on the NewOnly Zealand insects, and the effects on 19 20 different lineages are discussed. The New Zealand insects are unique, yet many are threatened 21 22 with extinction and efforts to preserve the fauna are reviewed. Despite the accumulating 23 24 knowledge, major gaps still exist and these are outlined, as are opportunities to address key 25 26 questions. The review ends with a synthesis and a discussion of how systematics, new 27 technologies, and integrative approaches have the promise to improve dramatically our 28 29 understanding of New Zealand insect evolution. 30 31 32 33 34 Key words biogeography, adaptation, alpine, freshwater, island 35 36 37 38 39 40 INTRODUCTION 41 42 43 44 45 The origins and evolution of the New Zealand biota have been of great interest to biologists ever 46 47 since Owen (1843) began describing moa bones, and Darwin’s (1859) seminal work. Although 48 well known for the relictual lineages within the vertebrate fauna (Tennyson 2010; Allentoft & 49 50 Rawlence 2012; Mitchell et al . 2014; Rheindt et al . 2014), the New Zealand insect fauna is 51 52 equally as interesting (Fig 1). This fauna also contains relictual lineages (Leschen 2006), as well 53 54 as unusual radiations (Fleming 1975; Marshall et al . 2008), uncharacteristically depauperate 55 56 lineages (Gibbs 1980; Watt 1975), gigantism (Kuschel & Worthy 1996; Leschen & Rhode 2002; 57 Gibbs 1998; Painting et al . 2014), and many unusual ecological interactions (Dugdale 1975). In 58 59 60 Page 3 of 81 Austral Entomology

3 1 2 3 addition to understanding the evolution of particular lineages and taxa that display the above 4 5 features, the New Zealand insects offer unlimited opportunities for deepening our understanding 6 7 of fundamental evolutionary processes such as speciation (Marshall et al . 2008, 2011; Albre et 8 9 al . 2012), adaptation (Buckley & Simon 2007; McCulloch et al . 2009; Dunning et al . 2013a, 10 11 2014), hybridisation (e.g. Chinn & Gemmell 2004; MorganRichards & Wallis 2003; Morgan 12 Richards et al . 2010), and phylogeography (e.g. Marske et al . 2012; Goldberg & Trewick 2011). 13 14 Previously, much research on the evolution of New Zealand insects has focussed on pattern 15 16 based questions, such as the enduring vicariance versus dispersal debate (e.g. Craw 1988; 17 18 Trewick & Wallis 2001).For However, Review many other questio nsOnly and conundrums have been neglected, 19 which we highlight and discuss here. Biogeographic patterns in the New Zealand insect fauna 20 21 have been reviewed comprehensively at broad (e.g. Goldberg et al . 2008; Trewick et al. 2011) 22 23 and more local scales (e.g. Buckley & Leschen 2013) and we do not attempt to update these 24 25 reviews here. Our purpose is to cover outstanding questions on the evolution of the New Zealand 26 insect fauna, to summarise current knowledge, and then to outline a prospective research agenda 27 28 designed to advance the field of the evolutionary biology of New Zealand insects. Although this 29 30 review is limited to the insects, we mention other terrestrial and aquatic invertebrate taxa where 31 32 they illustrate a pattern or process that has been poorly studied in the insects, yet likely to be of 33 34 importance. 35 36 We divide this examination of the New Zealand insect fauna into several sections. The 37 38 first four sections review the origins and evolution of four different components of the insect 39 40 fauna; terrestrial insects from lowland habitats, alpine insects, freshwater insects, and insect 41 42 communities on New Zealand offshore islands. The debate on the origins of the fauna has been 43 at the heart of New Zealand insect biogeography for many years and much recent progress has 44 45 been made with phylogenetic approaches. Nevertheless, large elements of the insect biota, 46 47 particularly from the cryptic and hyperdiverse soil and saprophagous communities, have barely 48 49 been examined. 50 51 We discuss landscape changes that occurred during the Cenozoic and in particular the 52 53 reduction in land area during the Late Oligocene, postulated to have caused massive turnover in 54 55 terrestrial biodiversity (e.g. Cooper & Cooper 1995; Landis et al . 2008). Moreover, New Zealand 56 57 harbours diverse terrestrial ecosystems with numerous fascinating species interactions that are as 58 59 60 Austral Entomology Page 4 of 81

4 1 2 3 complex and intricate as on any landmass. We mention several systems and focus in particular 4 5 on the sooty mould biota, a complex web of interactions between insects, fungi and plants. The 6 7 freshwater and alpine zone insect faunas are well known for their endemism and unusual 8 9 adaptations and we discuss the origins of this diversity. We focus particularly on the unusual 10 11 physiological and biochemical adaptations that have enabled insect lineages to radiate into alpine 12 environments. These adaptations include suites of behavioural, physiological, and biochemical 13 14 strategies and we are only beginning to the scratch the surface of the underlying evolutionary 15 16 processes (Sinclair et al . 1999; Dunning et al . 2014). In a following section, we assess the 17 18 influence of PleistoceneFor glaciation Review and climate change in Onlyshaping the diversity and distributions 19 of New Zealand insects. We review the effects these processes have had on the geographic 20 21 distributions of lineages at both the interspecific and intraspecific levels and discuss evidence for 22 23 the geographic location and nature of glacial refugia. We also address the insect fauna of New 24 25 Zealand offshore islands as these areas are a key focus for conservation biologists as refuges for 26 wildlife that has been driven to extinction on the mainland (Towns et al . 2012). Furthermore, 27 28 these islands contain many endemic lineages that are a result of the individual geological 29 30 histories of each island (e.g. Buckley & Leschen 2013). We summarise the phylogenetic patterns 31 32 and discuss some unique features of the island insect faunas, relating these to the history of land 33 34 connections to the New Zealand mainland. We discuss the current conservation threats and 35 requirements of the New Zealand insect fauna that are underway to ensure this unique 36 37 biodiversity survives for future generations to enjoy. This review concludes with a synthesis and 38 39 a summary of promising future research directions. 40 41 42 43 44 AGE AND ORIGINS OF NEW ZEALAND TERRESTRIAL INSECTS 45 46 47 48 49 How old is the terrestrial insect fauna and what are its origins? 50 51 52 53 54 55 The New Zealand terrestrial insect fauna is characterised by its disharmonic nature and mixture 56 of old and young lineages (Watt 1975). Although levels of endemism at the generic and species 57 58 59 60 Page 5 of 81 Austral Entomology

5 1 2 3 level are very high (Watt 1975; Gibbs 2006), there are only six insect families that are endemic 4 5 namely: Mnesarchaeidae moths (Gibbs 1989), Mystacinobiidae bat flies (Holloway 1976), and 6 7 the four beetle families Agapythidae (Leschen et al . 2005), Chalcodryidae (Watt 1974), 8 9 Metaxinidae (Kolibáč 2004), and Cyclaxyridae (Gimmel et al . 2009). Some insect lineages that 10 11 are highly diverse in neighbouring landmasses are either absent in New Zealand, such as many 12 beetle families, or have relatively low diversity, such as Orthoptera, Neuroptera, Apoidea, 13 14 Formicidae, and the butterflies (Watt 1975; Gibbs 1980). Other lineages are highly diverse in 15 16 New Zealand, yet typically have lower relative diversity elsewhere, such as Zopheridae beetles 17 18 (Lord & Leschen 2014)For and Oecophoridae Review moths (Hoare Only2005, 2010). This disharmony may be 19 related to the longterm isolation of the New Zealand landmass and absence of other key groups 20 21 leading to novel ecological opportunities (Duthie et al . 2006) and dramatic environmental 22 23 changes causing unequal rates of species turnover among lineages (e.g. Watt 1975). 24 25 Understanding this disharmony and the composite nature of old and young lineages requires an 26 understanding of the timing and processes by which insect lineages have arrived in New Zealand 27 28 and subsequently evolved and diversified. 29 30 31 Biogeographic studies to determine the geographic location of sister taxa to New Zealand 32 33 clades and then infer the process by which these lineages have come to inhabit New Zealand 34 have revealed several patterns. New Caledonia was the Gondwanan landmass to which New 35 36 Zealand was connected most recently (Herzer et al ., 1997; Lee et al . 2001; Ladiges & Cantrill, 37 38 2007; Schellart et al ., 2009), and not unexpectedly some New Zealand insect lineages have sister 39 40 taxa in New Caledonia, including cicadas (Arensburger et al . 2004), stick insects (Buckley et al . 41 42 2009a, 2010a), dung beetles (Monaghan et al . 2007), and possibly some of the Anostostomatidae 43 weta (Pratt et al . 2008). However, many questions remain unanswered about the nature of this 44 45 biogeographic connection. One significant question is the uncertainty around the timing of the 46 47 final breaking of land connections between New Zealand, New Caledonia, and Australia, which 48 49 may not have occurred until 55 million years ago (Schellart et al . 2006; Allentoft & Rawlence 50 2012) or even later (Ladiges & Cantrill 2007). Many authors interpret divergence dates between 51 52 New Zealand lineages and their sister taxa in Australia or New Caledonia of much less than 80 53 54 million years ago as indicating dispersal rather than a vicariant origin. However, if these latter 55 56 dates for the final rupturing of land connections to the north of New Zealand are correct then 57 58 some previous conclusions about dispersal need revisiting. The movement of island arcs, 59 60 Austral Entomology Page 6 of 81

6 1 2 3 especially on the Norfolk, Three Kings and Reinga Ridges in the Cenozoic could have acted to 4 5 transport biota from New Caledonia to New Zealand (Herzer et al . 2007; Buckley et al . 2010a). 6 7 Schellart et al . (2009) also argued for an arc of volcanoes between New Caledonia and New 8 9 Zealand along the Loyalty and Three Kings Ridges during the Cenozoic that could have provided 10 11 a means for island hopping at least between the two landmasses. Undoubtedly many lineages 12 have arrived by longdistance dispersal (e.g. Arensburger et al . 2004). Others studies have 13 14 inferred sister group relationships between New Zealand and Australia (e.g. Arensburger et al. 15 16 2004; Allegrucci et al . 2010; Kayaalp et al. 2013), indicative of a range of processes leading to 17 18 the assembly of the insectFor fauna. TheReview geology of the Southwest Only Pacific is certainly complex, yet 19 more detailed reconstructions of the history of emergent land since the late Cretaceous are 20 21 required to gain a better understanding of the mechanisms underlying sister taxa relationships 22 23 between New Zealand and New Caledonia. 24 25 26 The absence of some insect groups in New Zealand, when they are present in 27 neighbouring landmasses, can be attributed either to; 1) the lineage being younger than the 28 29 rifting date, 2) the lineage being older than the rifting date, yet absent in the New Zealand region 30 31 of Gondwana prior to rifting, and 3) a post rifting extinction in New Zealand (Craw & Waters 32 33 2006; Giribet & Boyer 2010; Sharma & Wheeler 2013). The lack of clarity around the timing of 34 the loss of land connections makes it difficult to determine if absence of a lineage is due to it 35 36 being younger than the age of rifting. Undoubtedly biotic regionalisation existed across 37 38 Gondwana (e.g., Scotese 2004; Murienne et al . 2014) and this may have influenced the absence 39 40 of some lineages from modern New Zealand. Post rifting extinction clearly has played a role in 41 42 the evolution of the flora, with many plant genera present in the fossil record absent in the 43 modern flora (e.g. Lee et al . 2001). However, with such a poor fossil insect record (e.g. Kaulfuss 44 45 et al . 2010), extinctions are very difficult to detect in insects. The loss of major plant groups such 46 47 as Eucalyptus , Acacia , Casuarina , and many Proteaceae species since the Miocene likely would 48 49 have caused extinctions in the insect fauna among specialised herbivores. Furthermore, host 50 shifts would have occurred as herbivores adapted to new host plants when their older hosts 51 52 became extinct. For example, Australian congeners of the New Zealand cecidomyiid fly 53 54 Eucalyptodiplosis chionochloae Kolesik, Safarti, Brockerhoff and Kelly form galls exclusively 55 56 on Eucalyptus . Yet in New Zealand E. chionochloae feeds on seeds from the tussock grasses 57 58 Chionochloa (Kolesik et al . 2007). Possibly this host shift is the result either of extinction of the 59 60 Page 7 of 81 Austral Entomology

7 1 2 3 previous Eucalyptus host in New Zealand, or alternatively dispersal from Australia following the 4 5 extinction for Eucalyptus and an immediate host switch to Chionochloa . Floral turnovers may 6 7 have also influenced specialised saproxylic beetles that would have tracked changes in coarse 8 9 woody debris. Other species that are polyphagous on their deadwood hosts and/or fungi may 10 11 have been less strongly effected (Leschen 2006), but further research is required. 12 13 14 15 16 The Oligocene Drowning and changing shorelines 17 18 For Review Only 19 20 21 Following the rifting from the eastern margin of Gondwana, New Zealand’s land area began to 22 23 erode during the Paleocene and by the late Oligocene had decreased to approximately 15% of its 24 25 current land area (Fleming 1979; Cooper & Cooper 1995; Landis et al . 2008; Scott et al . 2014). 26 The “Oligocene Drowning” hypothesis of Cooper and Cooper (1995) predicted that this 27 28 reduction in land area caused a bottleneck in vertebrate lineages that can be observed in the post 29 30 Oligocene radiations of some vertebrate groups (Cooper & Cooper 1995; Bunce et al . 2009). 31 32 However, the effect of the Oligocene Drowning on the insect fauna is still very poorly known. 33 Although the exact area of land during the maximum extent of the drowning has not been 34 35 determined (Scott et al . 2014), Lee et al . (2001) argued that the total land area would have 36 37 exceeded that of modern New Caledonia, which itself harbours a diverse insect fauna (Chazeau 38 39 1993). This raises the prospect that the Oligocene Drowning may have had little effect on the 40 41 diversity of insects, unlike the vertebrate fauna, which could have been more susceptible to a 42 reduction in land area. However, if the total land area was in fact less than that proposed by 43 44 Cooper and Cooper (1995) and Lee et al . (2001) then the impact on the insect fauna may have 45 46 been more significant. Resolution of this issue will require dated phylogenies for a range of 47 48 lineages distributed across the New Zealand region with appropriate outgroups, that survived 49 through the Oligocene. Selection of lineages that are younger than the breakup of Gondwana 50 51 leads to weak tests of vicariance as pointed out by Murienne et al . (2014), but application of 52 53 diversification models (e.g. Rabosky 2006) to these phylogenies will enable robust testing of 54 55 these hypotheses. If a bottleneck can be observed in New Zealand insect lineages then this begs 56 57 the question as to what are the intrinsic and extrinsic factors that enabled some lineages to 58 59 60 Austral Entomology Page 8 of 81

8 1 2 3 survive with their diversity intact whereas other lineages experienced elevated extinction rates. 4 5 The application of diversification models with dated phylogenies and species traits will assist in 6 7 resolving this question. 8 9 10 Recent geological models for the Cenozoic evolution of the New Zealand landmass have 11 shown a 500 km displacement along the alpine fault, most of which has occurred over the past 23 12 13 million years (Bunce et al . 2009). The associated shoreline reconstructions show a lack of direct 14 15 land connections between northern and southern New Zealand from the early Miocene until the 16 17 Pleistocene (Bunce et al . 2009; Trewick & Bland 2012). This reconstruction suggests the 18 Cenozoic geological historyFor of New Review Zealand is defined byOnly a “northern” and a “southern” island 19 20 or island groups that do not correspond exactly to the modern North and South Islands. These 21 22 models show a seaway between the “southern” and “northern” islands since the Miocene that has 23 24 periodically had smaller islands within it. The general geological pattern is that of a southern 25 26 migration of this seaway. Furthermore, the southern tip of the Wellington region had land 27 connections with the upper South Island throughout the Miocene and Pliocene to as recently as 28 29 less than 1 million years ago when the modern Cook Strait formed (Bunce et al . 2009). This 30 31 model has significant implications for the interpretation of biogeographic features and 32 33 phylogenetic patterns across New Zealand (e.g. Stevens & Hogg 2004; Buckley et al . 2011; 34 Trewick & Bland 2012). For example, it has long been noted that there is a biogeographic and 35 36 phylogeographic break across the middle of the North Island (see Taupo Line, below) and this 37 38 could be explained in part by the Cenozoic “northern” and “southern” islands. Furthermore, 39 40 many species and lineages are distributed from the South Island across Cook Strait into the lower 41 42 North Island and this can potentially be explained by the Cenozoic South Island – lower North 43 Island land connection (e.g. Marra et al . 2008; Marske et al . 2011; Trewick & Bland 2012). The 44 45 ephemeral islands in the Cenozoic seaway may also be responsible for the number of endemic 46 47 insect species and lineages found in southern Hawke’s Bay area (e.g. MorganRichards & Gibbs 48 49 1995; Marshall et al . 2011). 50 51 52 53 Ecological relationships of terrestrial insects 54 55 56 57 58 59 60 Page 9 of 81 Austral Entomology

9 1 2 3 The disharmonic nature of the New Zealand biota includes occupancy by many species of 4 5 atypical or novel niches (e.g. Duthie et al . 2006) and the formation of unusual ecological 6 7 interactions and cospeciation events (e.g. Dugdale 1975). Hostplant interactions in New Zealand 8 9 have not been covered broadly, though specific relationships exist in many groups of herbivore 10 11 lineages, especially weevils (e.g. Kuschel 1971, 1990; May 1993) and Lepidoptera (Dugdale 12 1975). Also poorly known are insecthost fungal relationships, with an unusual association 13 14 between many species of beetles and various taxa of polyporoid and corticioid fungi (e.g. Marske 15 16 et al . 2009; Yavorskya et al . 2014). In this case, fungal polyphagy is predominant in 17 18 mycophagous insects, Forwith few associations Review of monophagy Only like the very specific host 19 associations of beetles on Ganoderma fungi (e.g. Kadowaki et al . 2011a,b). This latter 20 21 interaction also includes one corylophid that has a larva with a weevillike snout for feeding on 22 23 spores deeply within the hymenium (i.e. Kadowaki et al . 2011b; Yavorskya et al . 2014). 24 25 26 A notable fungal relationship is the sooty mould fauna; notable because sooty moulds can 27 form huge masses on several tree species, mainly Nothofagaceae and Leptospermum , and 28 29 accumulate on any stationary surface. Sooty moulds are found worldwide (Chomnunti et al . 30 31 2014) but are particularly abundant in New Zealand (Hughes 1976). These fungi thrive 32 33 especially on the exudates of stenorrhynchan Hemiptera and in New Zealand large masses form 34 on the exudates of the scale insect Ultracoelostoma assimile (Maskell) (Morales et al . 1988). 35 36 Most extraordinary are their thick masses in beech forests in the South Island, especially in 37 38 Canterbury and Northwest Nelson. These host many species of insects (Morales et al . 1988), 39 40 especially beetles, and include derodontids, nitidulids, salpingids, anthribids (e.g., Klimaszewski 41 42 & Watt 1997; Carlton & Leschen 2007) and the endemic families Metaxinidae (Kolibáč 2004; 43 Johnson et al. 2008; 1 sp.), Agapythidae (Leschen et al. 1995; 1 sp.), and Cyclaxyridae (Gimmel 44 45 et al . 2009; 2 spp.). The exact number of true sooty mould specialists is unknown because 46 47 detailed surveys have not been conducted throughout New Zealand. Moreover, exact diets of the 48 49 sooty mould beetles are unclear. Some species may be feeding as parasites on the scale insects 50 (Morales et al . 1988) and others feeding on the sooty moulds and/or the exudates as reported in 51 52 other predators of hemipterans (Leschen 2000). 53 54 55 The unusual number of endemic beetle families associated with sooty moulds indicates 56 57 an ancient association and this is confirmed by recent molecular evidence that shows 58 59 60 Austral Entomology Page 10 of 81

10 1 2 3 Cyclaxyridae and Agapythidae diverged from their sister taxa in the Cretaceous, Metaxinidae in 4 5 the Paleogene, while Nothoderodontus Crowson (a derodontid genus distributed also in Chile 6 7 and Australia) is dated to the Jurassic (Duane McKenna, personal communication). If the 8 9 remaining members of the sooty mould fauna are as old as these examples, future systematics 10 11 studies of groups that contain a mix of sooty mould and “freeliving” species should reveal older 12 sooty mould lineages relative to sister taxa. 13 14 15 Meanwhile, a few specialist groups of which are present in New Zealand, that occur in 16 17 comparatively higher numbers elsewhere (see freshwater commensal midges below). For 18 example, there are fewFor species of Revieweusocial insects with only Only eleven species of native ants (Don 19 20 2007) and one termite ( Kalotermes brouni Frogatt), though one extinct species has been recently 21 22 described from the Miocene (Kaulfuss et al . 2010). Ant and termite inquilines are common 23 24 throughout the world but there is only one species of New Zealand beetle inquiline repeatedly 25 26 reported in the literature: Holloceratognathus passaliformis (Holloway) (Lucanidae) described 27 from nests of Prolasius advena (Fr. Smith) (Holloway 2007). The lack of inquilines, however, is 28 29 due to bias in collecting and lack of detailed studies, as there are several staphylinid beetles 30 31 associated with three species of native ant (S. Nomura and R.A.B. Leschen, in prep.). The 32 33 preceding inquiline and sooty mould examples and other communitylevel phenomena, like the 34 overdominance of white flowers (Heine 1937; Primack 1978, 1993; Swenson & Bremer 1997), 35 36 indicate that many of New Zealand’s ecological relationships require detailed study, or have yet 37 38 to be discovered or appreciated in the scientific literature. 39 40 41 42 43 ORIGINS AND SPECIATION OF ALPINE INSECTS 44 45 46 47 48 49 Age of the alpine environment and the insect fauna 50 51 52 53 54 A dominant geographic feature of New Zealand are the main axial mountain ranges that run the 55 56 length of South Island and into the southern North Island. They reach their peak in the central 57 region of the Southern Alps where Aoraki / Mount Cook is 3,764 metres in elevation and well 58 59 60 Page 11 of 81 Austral Entomology

11 1 2 3 above the summer snow line. The rate of uplift of the Southern Alps as well as other ranges such 4 5 as the Kaikoura Range is particularly rapid by international standards (Tippett & Kamp 1993; 6 7 Kao 2001; Robinson et al . 2004). This, coupled with the relatively soft sedimentary rock of 8 9 which many mountain ranges are comprised and extensive faulting, has led to a high erosion rate 10 11 and unstable landscape. The landscape has been further influenced by the erosional effects of 12 recent glaciation. The mountains of North Island are by contrast much lower than the Southern 13 14 Alps, with the high point being the active volcano Mount Ruapehu at 2,797 metres, below the 15 16 summer snow line. The other alpine zones of North Island are fewer and much smaller in size, 17 18 scattered and often separatedFor by largeReview distances in contrast Only to South Island. 19 20 The New Zealand alpine flora is highly endemic at the species level (McGlone et al . 21 22 2001) and similarly for the insect fauna, although this has yet to be quantified. There are also 23 24 strong biogeographic patterns with many groups, such as the cicada genus Maoricicada Dugdale 25 26 showing higher species richness in the northern and southern regions of South Island relative to 27 the central South Island and North Island (Dugdale & Fleming 1978). Chinn and Gemmell 28 29 (2004) demonstrated that cockroaches of the genus Celatoblatta Johns share mitochondrial 30 31 haplotypes in the central region of the Southern Alps, yet were monophyletic at this locus in the 32 33 northern and southern alpine areas of South Island. This was attributed to secondary contact 34 between species previously isolated to the north and south of the Southern Alps due to the high 35 36 rate of uplift, erosion and other environmental changes in the central region of South Island. 37 38 Similar processes are likely occurring in other alpine insect groups such as grasshoppers 39 40 (Trewick 2007). Some alpine species also show high levels of genetic variation among 41 42 populations, with little or no sharing of haplotypes (e.g. Trewick et al . 2001; O’Neill et al . 2009; 43 Dunning et al . 2014), indicative of reduced gene flow between alpine populations isolated on 44 45 “sky islands”. At the species level some species are widespread across large areas (e.g. 46 47 Maoricicada cicadas; Dugdale & Fleming 1978) whereas others are restricted to single mountain 48 49 ranges (e.g. Syrphetodes Pascoe beetles; Leschen & Buckley 2014). Identification of these areas 50 or local endemism, disconnected from surrounding populations, will be critical for species 51 52 management as geographic distributions shift due to climate change. Although there are clear 53 54 biogeographic patterns across the New Zealand alpine zone, we lack enough phylogenetic 55 56 reconstructions from diverse insect groups to identify common driving forces. 57 58 59 60 Austral Entomology Page 12 of 81

12 1 2 3 A remarkable aspect of the New Zealand alpine habitat is the extraordinary diversity of 4 5 plants and animals coupled with its young age (Heenan & McGlone 2012). Permanent and 6 7 widespread high alpine habitats above the tree line became established only <1.5 million years 8 9 ago (mya), although upland and open habitats have existed since the Miocene. Heenan and 10 11 McGlone (2012) reviewed the ages of plant lineages that inhabit various parts of the alpine 12 environment. Boginhabiting lineages were found to be older than those of open habitat 13 14 generalists, which in turn were older than the alpine specialists, consistent with alpine specialists 15 16 having evolved from ancestors inhabiting open habitats. The alpine specialists for which 17 18 molecular dates are availableFor were Review all less than 2 million Onlyyears old, consistent with the first 19 appearance of a permanent alpine zone (< 1.5 mya). Buckley and Simon (2007) reviewed the 20 21 ages of lineages that contained species found in alpine habitats. These ages ranged from 2.6 mya 22 23 for the grasshopper genus Alpinacris Bigelow (Trewick & Wallis 2001) to 17 million years for 24 25 the weta genera Hemideina Walker and Deinacrida White (Trewick & MorganRichards 2005). 26 These dates are overestimates for the origin of the alpine species because they are crown 27 28 diversification ages for clades containing both lowland and alpine species. For example, Buckley 29 30 and Simon (2007) estimated a crown age for Maoricicada from the mid to late Miocene. 31 32 However, this genus contains lowaltitude open habitat species in addition to high alpine 33 34 specialists. The diversification of the high alpine Maoricicada specialists was estimated to have 35 occurred in the Pliocene and Pleistocene, in agreement with Heenan and McGlone’s (2012) 36 37 model. Furthermore, Buckley and Simon (2007) estimated a low elevation habitat for the 38 39 ancestor of the alpine species using ancestral character state reconstruction, again consistent with 40 41 Heenan and McGlone (2012). Although this model has been validated for several insect lineages 42 further studies are required for taxa encompassing a broad range of habitats, with well dated 43 44 nodes at the point that habitat transitions have occurred. 45 46 47 As noted above, alpine insect species richness varies across New Zealand. Although 48 49 figures have not been compiled for the alpine insects, like they have for vascular plants (e.g. 50 Wardle 1963, 1988), similar patterns have become apparent in some insect groups (e.g. Dugdale 51 52 & Fleming 1978). Notable patterns include the low levels of endemism on the volcanic uplands 53 54 of North Island, and high levels of endemism in the vast alpine expanse of Northwest Nelson. 55 56 More detailed information is required on alpine insect species distributions across New Zealand 57 58 and this will require intensive and specialised sampling of inaccessible areas such as Fiordland 59 60 Page 13 of 81 Austral Entomology

13 1 2 3 and Northwest Nelson. Only when such information is available will we be able to reconcile 4 5 insect and higher vascular plant distribution patterns. 6 7 8 9 10 Adaptation to the alpine environment 11 12 13 14 15 16 The alpine zone is inhabited by all insect orders present elsewhere in New Zealand and these 17 lineages have acquired the adaptations required for alpine survival in a remarkably short time. 18 For Review Only 19 Insect species that colonise these environments contend with a variety of challenges including; 20 21 lower temperatures with a higher frequency of freezing events than lower altitudes, cold 22 23 associated desiccation, diet shifts due to highly modified host plants, and shortened seasons for 24 25 juvenile growth and adult reproduction (Sinclair et al . 2003b; Wharton 2011; Dennis et al . 26 2014). Many New Zealand insect lineages evidently have adapted to these challenges through 27 28 modification of their behavioural, morphological, physiological, and biochemical phenotypes 29 30 (Wharton 2011). 31 32 33 Some of the more notable alpine lineages include the beetle genus Syrphetodes , a 34 member of the Ulodidae found across Australia, New Caledonia, and South America that always 35 36 inhabits forest habitats where the larvae feed on dead wood. However, alpine species in New 37 38 Zealand are often found far above the tree line in habitats devoid of woody vegetation; clearly 39 40 there has been a unique and dramatic habitat and host shift (Leschen & Buckley 2014). Many 41 alpine insects are notable for their dark colour relative to lowland sister taxa, and this 42 43 phenomenon has been observed in the cicada genus Maoricicada (Fleming 1975; Buckley & 44 45 Simon 2007), the terrestrial stonefly Holcoperla McLellan (McLellan 2006; McCulloch et al . 46 47 2010), Syrphetodes beetles (Leschen & Buckley 2014), Percnodaimon Butler butterflies (Gibbs 48 49 1980) and the alpine grasshopper Brachaspis robustus Bigelow (Trewick 2001). Many of these 50 same insects demonstrate behavioural adaptations such as basking (e.g., Gibbs 1980). One 51 52 notable adaptation that appears to be rare in the insect fauna as a whole is diapause, which is 53 54 common in Northern Hemisphere insects (Roberts 1978; Morris 1989). Instead, many insects 55 56 show quiescence rather than full diapause during winter months in cooler areas, which has been 57 58 59 60 Austral Entomology Page 14 of 81

14 1 2 3 attributed to the milder climate regime in New Zealand relative to the continental Northern 4 5 Hemisphere (Dumbleton 1967; Morris 1989). 6 7 8 Although little is known about these behavioural adaptations, even less is known about 9 10 the physiological, metabolic, and biochemical adaptations that have allowed these alpine insects 11 to radiate into the alpine environment (Sinclair et al . 1999). Alpine taxa in some insect clades 12 13 form monophyletic groups to the exclusion of lowland species (i.e. the alpine Syrphetodes 14 15 clade); however, in other insect lineages that include multiple alpine species these alpine taxa do 16 17 not form monophyletic groups (e.g. Emerson & Wallis 1995; MorganRichards & Gibbs 2001; 18 Dunning et al . 2013a).For These phylogenetic Review patterns raise Onlythe possibility of multiple, independent 19 20 colonizations of the alpine zone. We do not know if different, yet closely related lineages that 21 22 have radiated independently into the alpine zone use similar physiological and biochemical 23 24 strategies to survive this harsh environment (Wharton 2011). A recent study on the variation of 25 26 gene expression among populations and species of the stick insect genus Micrarchus Carl 27 (Dunning et al . 2014) showed large differences in gene expression among lowland and alpine 28 29 species and among populations of a single alpine species, unrelated to shared ancestry. This 30 31 suggests different stick insect species are using different strategies to survive in the alpine 32 33 environment. 34 35 Wharton (2011) recently reviewed current knowledge on the biochemical and 36 37 physiological basis of cold tolerance in New Zealand insects. Cold tolerance typically falls into 38 39 two categories: freeze tolerance and freeze avoidance. The former category is when the animal 40 41 can survive the formation of internal ice, and often controls where in the body and at what rate 42 ice forms. The latter is when the animal avoids the formation of internal ice through various 43 44 biochemical processes. In the Southern Hemisphere, freeze tolerance strategies seem to be more 45 46 common than freeze avoidance strategies, the reverse of the pattern observed in the Northern 47 48 Hemisphere (Sinclair et al . 2003a). This has been attributed to the milder yet variable climates of 49 50 the Southern Hemisphere (Sinclair et al . 2003b), though Holarctic lineages are more widespread 51 and may have undergone more frequent interchanges than Gondwanan groups (Sanmartín et al . 52 53 2001). Studies of the alpine weta Hemideina maori (Pictet & Saussure) (Ramløv 1992; Sinclair 54 55 et al . 1999) and the alpine cockroach Celatoblatta quinquemaculata Johns (Sinclair 1997) show 56 57 these are freeze tolerant, utilising ice nucleators, which control the temperature and location at 58 59 60 Page 15 of 81 Austral Entomology

15 1 2 3 which ice forms within the body. The gene expression response of one species of alpine stick 4 5 insect, Micrarchus sp. nov. 2, has been investigated using messenger RNA sequencing and real 6 7 time PCR (Dunning et al . 2013b; 2014). These studies showed that genes associated with the 8 9 cuticle tended to be expressed differentially relative to a lowland species, and thus modification 10 11 of the stick insect cuticle may protect this species against ice formation across the cuticle or 12 increasing desiccation resistance. 13 14 Both the weta and the cockroach are in clades that contain other alpine and lowland 15 16 species. With the exception of some data on supercooling points and ice formation in 17 18 haemolymph in two otherFor Hemideina Review species, in both cases Only very little is known about the cold 19 tolerance processes in related alpine and lowland species. Although we have information on the 20 21 basis of adaptation to the alpine environment in these species, what is required are detailed 22 23 studies on the comparative physiology, biochemistry and molecular genetics of cold tolerance 24 25 within radiations of species restricted to both the lowlands and alpine environments. Lineages 26 such as the cicadas (Buckley & Simon 2007), cockroaches (Chinn & Gemmell 2004), weta 27 28 (MorganRichards & Gibbs 2001; Trewick & MorganRichards 2005), stick insects (Buckley et 29 30 al . 2010a; Dunning et al . 2013a) and various beetle groups (e.g., Leschen & Buckley 2014) with 31 32 welldeveloped or emerging phylogenetic frameworks are ideal for advancing this goal. 33 34 35 36 37 THE EVOLUTION OF NEW ZEALAND FRESHWATER INSECTS 38 39 40 41 42 Whence (and when) did the freshwater fauna come? 43 44 45 46 47 48 The origins of New Zealand’s freshwater insects, like the terrestrial fauna, have long been a 49 contentious issue. Many workers have suggested that a large proportion of New Zealand’s 50 51 freshwater insect fauna are members of austral clades, with relatives in southeastern Australia 52 53 and South America, which represent Gondwanan relics (e.g. Brundin 1966; Winterbourn 1980; 54 55 Gibbs 2006; Craig et al . 2012). One significant question is the availability of freshwater habitats 56 during the Oligocene drowning (Cooper & Cooper 1995). Recently characterised fossil deposits 57 58 59 60 Austral Entomology Page 16 of 81

16 1 2 3 from the Eocene through Miocene suggest the presence of alluvial floodplain and lacustrine 4 5 habitats at various locations on the main islands (Ferguson et al. 2010; Lee et al. 2012). 6 7 Furthermore, late Oligoceneearly Miocene amber from sites on South Island has revealed adults 8 9 of numerous freshwater taxa (Kaulfuss et al. 2013), while compression fossils and their 10 11 depositional characteristics also imply the enduring presence of freshwater habitat from at least 12 the late Oligocene (Feldman & Pole 1994; McDowall & Pole 1997; Worthy et al. 2008; 13 14 Lindqvist & Lee 2009), and quite possibly much earlier (Feldman 1984). This suggests that not 15 16 only was there freshwater habitat present during this time, but also that it was inhabited by a 17 18 recognisable insect fauna.For Review Only 19 20 Gibbs (2006) suggested that extant New Zealand freshwater taxa are most likely to 21 22 represent Gondwanan relics because of the obvious restrictions on their successful dispersal 23 24 across lethal saltwater barriers and a general affinity for cold, pristine, lotic ecosystems. Recent 25 26 molecular phylogenetic studies of some freshwater insect groups also provide evidence for a 27 long history in New Zealand. For example, many groups within the New Zealand nonbiting 28 29 midges (Diptera: Chironomidae) appear to have undergone crown diversification in the mid 30 31 Eocene and follow expected area relationships for an austral ancestry (Cranston et al. 2010; 32 33 Krosch et al. 2011; Cranston et al . 2012; Krosch & Cranston 2013). New Zealand Plecoptera 34 diversification events range from the late Eocene to early Miocene and also largely follow an 35 36 austral pattern of relationships (McCulloch 2011). Moreover, it has been suggested that many 37 38 elements of the New Zealand freshwater fauna represent earlybranching lineages, thus inferred 39 40 (sometimes incorrectly) to be ‘primitive’ types reflective of a retained ancient fauna (Daugherty 41 42 et al . 1993; Collier 1993). For example, the petaltailed dragonflies (Family Petaluridae) were 43 long considered to be one such group, based on morphological characters, a supposition that has 44 45 recently been supported by molecular evidence (Ware et al . 2014). Taken together, it appears 46 47 very likely that there has been available freshwater habitat across the New Zealand landmass, 48 49 however much reduced during the Oligocene, since at least the Eocene and possibly even since 50 the initiation of rifting from Gondwana in the Cretaceous (~80 mya). 51 52 53 Colonisation of remnant or emergent habitats by newly arriving freshwater organisms is 54 55 likely to have been more complicated than for terrestrial taxa. For taxa that are restricted to 56 57 freshwater habitats for their entire life history, clearly their dispersal capabilities across saltwater 58 59 60 Page 17 of 81 Austral Entomology

17 1 2 3 barriers would have limited their colonisation of New Zealand. For taxa with an adult flight 4 5 stage, availability of appropriate habitat (lotic vs. lentic), adult stage duration, life cycle 6 7 seasonality, existing food resources and dominant resident taxa will also have had an impact on 8 9 the success of a colonising individual (Mackay 1992). Furthermore, dispersal in many flighted 10 11 freshwater insect taxa often is limited to the gravid female seeking an oviposition site. Thus, 12 successful colonisation of new habitats by such taxa requires the female to also find a suitable 13 14 site to lay her eggs and sufficient survival of the offspring to found a population. Immature 15 16 stages of most freshwater insects require large amounts of energy for growth and development, 17 18 and in some taxa suchFor as the Chironomidae Review this is the major Only feeding life stage. Most other insect 19 groups also feed as adults to fuel dispersal and mating activity and although there is significant 20 21 variation in life stage duration across the orders, immature stages are generally shorter than the 22 23 adult stage. At all life stages there is thus pressure on colonising individuals to locate food 24 25 resources and this may potentially favour generalist taxa as the most likely to successfully 26 establish in a new habitat. 27 28 29 Of those freshwater taxa inferred to have arrived in New Zealand after Gondwanan 30 31 fragmentation, many have putative sister groups in Australia or elsewhere in the south Pacific 32 33 region. Before the formation of the Antarctic Circumpolar Current and the West Wind Drift 34 phenomenon, aerial dispersal of freshwater insects may have been facilitated or at least aided by 35 36 other major meteorological events such as cyclone fronts. An alternative pathway, which may 37 38 also have aided fully aquatic taxa, may have been via a series of island chains along the Lord 39 40 Howe/Norfolk Island Rises that maintained connectivity between New Zealand and the 41 42 surrounding area during the Palaeogene (Veevers 2012; Krosch & Cranston 2013). Following 43 inception of West Wind Drift around 30 mya (Cook & Crisp 2005), asymmetric dispersal from 44 45 Australia to New Zealand would have been significantly more probable. Indeed, in more recent 46 47 times, a socalled ‘rain’ of invertebrates has been recorded dispersing from Australia to New 48 49 Zealand on wind currents (Tomlinson 1973; Close et al . 1978), including many freshwater taxa. 50 These abiotic phenomena, combined with intrinsic life history traits, clearly have influenced 51 52 which taxa have colonised New Zealand freshwater systems successfully. 53 54 55 56 57 Patterns of diversity in the freshwater taxa 58 59 60 Austral Entomology Page 18 of 81

18 1 2 3 4 5 6 Modern day New Zealand freshwater habitats range from lentic bogs and marshes, including 7 8 highaltitude tarns, to fastflowing seasonal glacierfed rivers, volcanic maar lakes to 9 10 groundwaterfed springs. There are heavily denuded, exposed stream channels, and those hidden 11 deep in prehistoric forest. Braided glacial rivers meander through steepsided valleys, forest 12 13 streams trickle down hillsides and waterfalls cascade from mountainsides. The wide ecological 14 15 breadth of New Zealand’s freshwater systems reflects a dynamic landmass in which widespread 16 17 volcanism, stochastic climatic patterns, glaciation and tectonic uplift all have played a role in 18 diversification of freshwaterFor habitat Review in New Zealand. In concert,Only these processes have been 19 20 central to the development and maintenance of a freshwater fauna very different to that found in 21 22 other temperate regions of the world, both at the community and species levels (Winterbourn et 23 24 al . 1981; Collier 1993). The dynamic nature of New Zealand’s climate and geology apparently 25 26 drove selection of taxa with wide environmental tolerances, life history flexibility and high 27 vagility (Winterbourn 1997). Many groups do not exhibit seasonal life history traits, such as 28 29 winter dormancy or diapause (see below), that are typical of taxa from temperate northern 30 31 hemisphere ecosystems (Dumbleton 1967; Winterbourn et al . 1981). This has been attributed to 32 33 both the presence of evergreen rather than deciduous trees that provide a yearround organic 34 energy source, and potentially that historical cold periods had little evolutionary influence on 35 36 New Zealand’s freshwater insects with regard to traits for cold avoidance (Dumbleton 1967). 37 38 Moreover, some functional groups (e.g. large particle detritivores) are underrepresented 39 40 compared with northern hemisphere temperate ecosystems (Winterbourn et al . 1981), mirroring 41 42 the ‘naturally depauperate’ patterns of the terrestrial fauna (Gibbs 2006). The lucky few groups 43 of taxa that either remained on the rifting landmass or arrived as new colonisers have since 44 45 diversified to fill niches occupied by other insect groups in temperate ecosystems elsewhere. A 46 47 result of this lowered diversity in many groups of New Zealand freshwater insects is that many 48 49 aquatic systems comprise similar suites of widespread taxa that are stable over time and despite 50 various forms of impact (Winterbourn 1997). Nevertheless, some freshwater insect groups do 51 52 exhibit significant specieslevel diversity and extreme endemism. Taken together, the New 53 54 Zealand freshwater fauna is enigmatic, with patterns of diversity and endemism variable across 55 56 many insect orders and taxonomic levels. 57 58 59 60 Page 19 of 81 Austral Entomology

19 1 2 3 Many freshwater insect orders, typically diverse elsewhere are not so in New Zealand and 4 5 yet exhibit high endemicity like the terrestrial fauna already mentioned. The Ephemeroptera are 6 7 represented by ~30 species (of ~220 in the Australasian region), all of which are endemic to New 8 9 Zealand, including two monospecific families (McLellan 1975; BarberJames et al . 2008). There 10 11 is a strong pattern of austral relationships among the New Zealand mayfly taxa, which supports a 12 potential Gondwanan origin (BarberJames et al . 2008). Odonata is only represented by 17 13 14 species (10 endemic to New Zealand), but this includes members of the family Petaluridae which 15 16 are supported as an ‘ancient’ group that predates New Zealand’s separation from Gondwana 17 18 (Rowe & Corbet 1987;For Daugherty Review et al . 1993; Kalkman etOnly al . 2008; Ware et al . 2014). Only six 19 freshwater Coleopteran families occur in New Zealand of ~25 known worldwide (Winterbourn 20 21 & Gregson 1981), including one endemic subfamily of Hydraenidae (Klimaszewski & Watt 22 23 1997). At least two species from the family Dytiscidae are known to inhabit cave freshwater 24 25 systems (Ordish 1976), ecosystems that are poorly characterised in New Zealand. Both 26 Megaloptera and Mecoptera are represented in New Zealand by only single species (Winterbourn 27 28 & Gregson 1981). The dobsonfly Archichauliodes diversus (Walker) is one of New Zealand’s 29 30 largest freshwater insects and inhabits clear, fastflowing streams on both main islands (Quinn & 31 32 Hickey 1990). The species can be quite abundant within streams and exhibits significant genetic 33 34 structure among populations (Hogg et al . 2002). The scorpionfly Microchorista philpotti 35 (Tillyard) has a highly restricted distribution and the poorly sclerotised larva occurs only in 36 37 protected slowmoving waters in streams in South Island. The taxon is thought to be closely 38 39 related to other austral members of the Nannochoristidae, potentially reflecting an ancient 40 41 Gondwanan heritage (Ferrington 2008). Only six families of aquatic/semiaquatic Hemiptera are 42 known, compared with 17 for the Australasian region (Winterbourn & Gregson 1981; Polhemus 43 44 & Polhemus 2008). Without taxonomically wellsampled, dated phylogenies it is difficult to 45 46 infer the origins and ancestry of many of these speciespoor New Zealand insect lineages, 47 48 particularly those that show affinities to taxa that inhabit similar ecosystems in other austral 49 50 regions. 51 52 By contrast, the Trichoptera are perhaps the best example of a group in which the New 53 54 Zealand fauna has diversified significantly in situ over at least the last ~23mya. The earliest 55 56 record of a caddisfly is a fossilised larval case from the Foulden Maar deposit, Otago, dated to 57 58 the earliest Miocene (U. Kaulfuss, unpub. data). Moreover, some groups currently restricted to 59 60 Austral Entomology Page 20 of 81

20 1 2 3 the austral regions (e.g. Helicophidae and psyllobetine Hydrobiosidae) are known from Eocene 4 5 Baltic amber (Botosaneanu & Wichard 1981), which suggests these groups may have been 6 7 retained after succumbing to extinction in the northern hemisphere (de Moor & Ivanov 2008). 8 9 The New Zealand fauna shows moderate endemicity at the genus level, but the order is highly 10 11 endemic at the species level. Some New Zealand taxa have developed high salinity tolerance and 12 indeed members of the endemic Chathamiidae family of marine caddisflies inhabit intertidal 13 14 zones throughout the New Zealand region. Within the transTasman family Conoesucidae, six 15 16 genera are endemic to New Zealand and their phylogenetic relationships suggest multiple 17 18 colonisation events (JohansonFor et alReview. 2009); however, no estimatesOnly of divergence dates currently 19 exist. The radiation of caddisfly taxa in New Zealand is thought by some to have been facilitated 20 21 by their holometabolous life history and diversity of available habitat for colonisation 22 23 (Winterbourn et al . 1981); however, this fails to explain the lack of diversification in insect 24 25 orders with similar life histories. 26 27 New Zealand Plecoptera have been particularly well characterised for species diversity, 28 29 with some 104 species known, 99 of which are from a single family (McLellan 2006). Taxa 30 31 included in McCulloch’s (2011) molecular phylogeny all exhibited connections with either 32 33 Australia or South America. Although New Zealand’s stoneflies generally prefer cool, flowing 34 freshwater systems (a trait generally associated with Gondwanan relic taxa), most New Zealand 35 36 stonefly groups are younger than Gondwanan age. Interestingly, it is thought that dispersal is 37 38 limited in stoneflies due to the ecological requirements of the nymphs and poor flight ability in 39 40 adults (Fochetti & de Figueroa 2008). It would seem unlikely, therefore, that these taxa would 41 42 have dispersed across wide ocean gaps to New Zealand, and so perhaps steppingstone island 43 connections were important for the movement of stoneflies also (McCulloch 2011). Furthermore, 44 45 numerous New Zealand genera have evolved to be brachypterous (reduced wings) or apterous 46 47 (wingless) as adults, often in combination with terrestrial nymphs (McLellan 2006). Wingless 48 49 New Zealand stoneflies generally are restricted to either the alpine zone of the major islands or 50 isolated habitats on some smaller offshore islands (McLellan 2006). Both life history states are 51 52 unusual for holometabolous freshwater taxa in an environmentally variable region like New 53 54 Zealand (Roff 1990). Typically, aptery is associated with relaxed selective pressures for dispersal 55 56 driven by environmental stability, as well as in specific habitats in which a flight stage may be 57 58 unnecessary. One possible explanation may thus be that the highly exposed and climatically 59 60 Page 21 of 81 Austral Entomology

21 1 2 3 variable nature of these ecosystems instead selected against dispersal in these taxa. In other 4 5 words, the evolutionary costs associated with flight in such environments (e.g. being blown off 6 7 course, high energy requirement) may have outweighed the benefits (e.g. increased gene flow 8 9 among populations), such that individuals that did not disperse possessed an evolutionary 10 11 advantage. 12 13 The Diptera are also considered to be diverse in New Zealand; however, there appears to 14 15 be marked variation in diversity across families. The Chironomidae are quite diverse with over 16 17 60 species known from five subfamilies (McLellen 1975), many of which represent austral 18 elements closely relatedFor to groups Review in Australia and South Only America, but with several other taxa 19 20 that represent more recent colonisations by cosmopolitan groups. The ecological breadth of this 21 22 group is similar to that found elsewhere, with species having radiated to inhabit most niches from 23 24 rapid glacial rivers to high altitude tarn pools to sealevel lakes. The species Chironomus 25 26 zealandicus Hudsonis perhaps the bestknown New Zealand chironomid as it is a pest of sewage 27 treatment works that can undergo such population explosion as to physically block pipeworks 28 29 (Forsyth 1971). At least three freshwater chironomids ( Tonnoirocladius commensalis Cranston, 30 31 Xenochironomus canterburyensis Forsyth and Eukiefferiella indet.) are commensals of 32 33 Blepheraceridae larvae, freshwater mussels and mayfly nymphs, respectively (Forsyth 1979; 34 Winterbourn 2004; Cranston 2007). Although commensalism is globally widespread in the 35 36 Chironomidae, this life history trait is rare among New Zealand freshwater insects as a whole. 37 38 Commensalism is thought to be an evolutionary strategy for maximising feeding opportunity, 39 40 mobility, and protection from disturbance while also reducing predation risk (Tokeshi 1993). 41 42 Certainly some environmental characteristics of New Zealand freshwater systems may indeed 43 have contributed to a selective advantage for commensalism in these taxa. Interestingly, T. 44 45 commensalis was placed phylogenetically within the coldadapted New Zealand genus Naonella 46 47 Boothroyd (Krosch et al . 2011), suggesting T. commensalis should be more accurately 48 49 designated as a species of Naonella and acts to highlight the lack of understanding surrounding 50 the development of such an unusual life history trait in these taxa. Another unusual chironomid, 51 52 the ‘iceworm’ midge Zelandochlus latipalpis Brundin, inhabits the Fox and Franz Josef Glaciers 53 54 on the west coast of South Island (Boothroyd & Cranston 1999). The larvae inhabit meltwater 55 56 pools and ice caves within the glacier and exhibit strong photophobia. Adult males inhabit the 57 58 glacier surface and are apterous, a trait that potentially developed under the same selection 59 60 Austral Entomology Page 22 of 81

22 1 2 3 pressures as argued for stoneflies above. A recent phylogenetic analysis suggests this taxon 4 5 diverged from its sister in the Oligocene (Cranston et al . 2010). Thus, the apparently high species 6 7 diversity and ecological breadth observed within the Chironomidae relative to other Dipteran 8 9 families may reflect a longer history in New Zealand and thus more time to radiate into different 10 11 niches. 12 13 Most of the remaining New Zealand Dipteran families do not possess such high species 14 15 level diversity and there is a mix of species endemic to New Zealand and those that are shared 16 17 with other landmasses in the south Pacific or more widely. Many of these groups are also 18 postulated to have an ancientFor vicariant Review origin on New Zealand Only (Tipulidae – de Jong et al . 2008; 19 20 Blepharaceridae – Wagner et al . 2008), although most have not been explicitly tested to date. 21 22 New Zealand also possesses a single representative of the Keroplatidae, the endemic glowworm 23 24 species Arachnocampa luminosa (Skuse), which inhabits caves and moist forest environments on 25 26 both main islands (Pugsley 1984). Arachnocampa occurs also in Australia, with A. luminosa 27 sister to a clade comprising all eight Australian species (Baker et al . 2008). The disjunct austral 28 29 distribution of the genus, along with the requirement for moist environments, poor adult 30 31 dispersal and extreme sensitivity to desiccation of these taxa raises questions concerning their 32 33 origin in New Zealand that have yet to be addressed adequately. 34 35 Overall, the New Zealand freshwater insect fauna comprises several groups of taxa that 36 37 are members of earlybranching lineages apparently retained from preGondwanan breakup 38 39 (e.g., petalurid dragonflies); a select few orders that presumably arrived after New Zealand 40 41 separated from Gondwana and have had a long history on the landmass, including surviving the 42 Oligocene inundation, such that they have diversified significantly in situ (e.g. stoneflies and 43 44 chironomids); and many groups that are species poor and either represent recently arrived poor 45 46 dispersers or ancient Gondwanan relic lineages that have been pruned by extinction. Thus, there 47 48 remain many unanswered questions concerning the evolution of New Zealand’s freshwater 49 50 insects, a conundrum that faces freshwater insects worldwide (Dijkstra et al. 2014). A substantial 51 proportion of New Zealand’s freshwater fauna are yet to be studied in a phylogenetic context and 52 53 determining the phylogenetic placement of New Zealand taxa amongst their austral relatives and 54 55 ages of diversification will contribute greatly to our understanding of the origins of these groups. 56 57 Even for studied taxa (e.g. the podonomine chironomids), knowledge gaps exist concerning the 58 59 60 Page 23 of 81 Austral Entomology

23 1 2 3 affinities of certain taxa amongst the broader New Zealand fauna. The search for fossil evidence 4 5 of freshwater insects will complement phylogenetic studies and if recent discoveries discussed 6 7 above are an indication, there could be a wealth of information yet to be revealed by the fossil 8 9 record. Moreover, incorporating ecological, physiological, and life history information in the 10 11 interpretation of resulting phylogenies will help to bring resolution to hypotheses about their 12 origins, as well as their diversification in New Zealand. 13 14 15 16 17 18 EVOLUTION OF INSECTSFor ON Review OFFSHORE ISLANDS Only 19 20 21 22 23 The New Zealand offshore islands 24 25 26 27 28 The geopolitical boundaries of New Zealand, extending from 34° north to 52° south latitudes, 29 30 encompass subtropical islands to Subantarctic Islands (Fig. 3). The Kermadec Islands, outside 31 32 this range, are absolutely tropical, and while they share many species in common with the 33 mainland, they also have a Pacific element (e.g. Brown et al . 2012). The distance from the New 34 35 Zealand mainland to these offshore islands varies, but in terms of diversity and conservation 36 37 value they are biological gems. The mostprotected nearshore islands (like HauturuoToi, 38 39 Kapiti Island, Takapourewa, and Codfish Island) are utilised by nesting pelagic birds and contain 40 41 species, mainly larger charismatic species of weevils and weta, that no longer exist on the 42 mainland due to introduced predators or loss of habitat. These islands also have endemic biota 43 44 whose ancestry provides insights into New Zealand’s dynamic biogeological history. There are 45 46 about 200 offshore islands, many under the stewardship of the Department of Conservation, 47 48 including the Subantarctic Islands which have World Heritage status. Only a handful of islands 49 have been intensely studied and typically not completely surveyed, and the most poorly studied 50 51 are those islands located in the complex network of fjords in Fiordland. We highlight the best 52 53 known of the offshore islands that are home to endemic invertebrates, have been moderately to 54 55 well surveyed, and are central for understanding the evolution of the New Zealand insect fauna. 56 57 58 59 60 Austral Entomology Page 24 of 81

24 1 2 3 The Northern Islands 4 5 6 7 8 9 The Three Kings Islands lie 50 km off the northern tip of North Island and can be easily seen on 10 a clear day. The largest island of the archipelago is Great Island or Manawatawhi (407 ha), 11 12 which is mostly cliffbound, the rest are mostly rock stacks and are all difficult to access. The 13 14 Three Kings Islands are an exposed part of the Three Kings Ridge, which is mostly submerged 15 16 and connected northward to the Norfolk Ridge, which is in turn joined to New Caledonia. The 17 fauna, in some respects, shows this connection (e.g. snails; Ponder et al . 2003), but absences, like 18 For Review Only 19 the erotylid beetle Kuschelengis Skelley and Leschen, known from the mainland and New 20 21 Caledonia, could be attributed to postrift extinction (see above). What is lacking, however, are 22 23 phylogenetic hypotheses for older groups present on the Three Kings, New Caledonia, New 24 25 Zealand, and elsewhere in the Pacific. Though visited several times by invertebrate researchers, a 26 full account of the species has not been published, but, based on unpublished work, 35% of the 27 28 beetle fauna may be endemic, including several endemic beetle genera (J. Marris, unpubl. data). 29 30 The great disparity between the mainland and the Three Kings also includes the range limits of 31 32 Pacific groups (snails), an AustralNeotropical beetle genus ( Platisus Erichson), and other 33 34 groups lacking immediate ancestors in New Zealand. Otherwise, the fauna is related to mainland 35 New Zealand, and in particular, to adjacent areas in Northland, while dated phylogenies 36 37 represent a range of divergences, with the oldest from the Miocene (Buckley & Leschen 2013). 38 39 40 Closer inshore to the Northland mainland at 22 km are the Poor Knights Islands, which 41 consist of Tawhiti Rahi (151.5 ha) and Aorangi (101 ha), a remnant that may have been 42 43 separated from the mainland at least during the Last Glacial Maximum (Hayward, 1991). Unlike 44 45 the Three Kings, there are no endemic plant species (the Poor Knights Lilly, Xeronema 46 47 callistemon , is found elsewhere [de Lange & Cameron 1999]), but it has been estimated that 16% 48 49 of the invertebrate fauna is endemic, with many species showing close relationships with species 50 found in Northland (Buckley & Leschen 2013), and in one case with the Three Kings Islands 51 52 (Seldon & Leschen 2011). Phylogenetic studies suggest regional endemism in the Northland 53 54 islands results from old diversifications associated with isolated islands that were oceanbound 55 56 during periods of higher sea levels (e.g. MorganRichards et al. 2001; Buckley et al . 2010b; 57 Leschen & Buckley 2014). Careful and dense regional sampling for phylogenetic studies will 58 59 60 Page 25 of 81 Austral Entomology

25 1 2 3 help distinguish divergence dates and determine if the ages of the island faunas relate to allopatry 4 5 following postglacial sea level rises (Buckley & Leschen 2013) or to preceding periods of land 6 7 reduction (Bunce et al . 2009). Also, the offshore islands, because of their range of distances 8 9 from the mainland, differences of island age, and area, represent an opportunity to test questions 10 11 of island biogeography. For example, we may be able to determine the differences between 12 palaeo and neoendemic species and the reason why mainland and islands populations tend to 13 14 express different phenotypes. For example, populations of the trogossitid beetle Phanodesta 15 16 brounii (Pascoe) on the Poor Knights Islands have aberrant elytral features (Leschen & Lackner 17 18 2013). For Review Only 19 20 21 22 23 The Chatham Islands 24 25 26 27 28 The Chatham Islands group is located on the Chatham Rise, is separated from the mainland by 29 30 about 680 km and is quite large (90,650 ha). Like the Three Kings Islands the Chathams Islands 31 32 has a high rate of beetle endemism of about 27%, with at least four endemic beetle genera 33 (Emberson 1998). This includes Thotmus halli Broun, collected from Pitt Island, known only 34 35 from the type specimen (Leschen et al . 2012) and thought to be extinct (Stringer & Hitchmough 36 37 2012). The Chatham Islands has a permanent residency of over 600 people, though there are 38 39 forest reserves and smaller islands that have intact faunas, and the smaller Pitt Island (6203), 40 41 which also supports a small population of people, has slightly more species (Emberson 1998). 42 Recent research (e.g., Trewick 2000) indicates that the biota has dispersed overwater since 2 mya 43 44 when it was presumably first exposed, despite dates much older for some taxa (Marske et al. 45 46 2012; Buckley & Leschen 2013), which may indicate potential sunken islands or continual land 47 48 elsewhere in the archipelago (Heenan et al . 2013). 49 50 Though questions regarding the source of the fauna remain unclear for some invertebrates 51 52 (Skelley & Leschen 2007), based on plant patterns, it is likely that sister groups are widespread 53 54 mainland taxa (Heenan et al . 2013). Dispersals and/or sharing ancestors from more distant areas, 55 56 like the scarab species Tesarius sulcipennis (Lea) described from Tasmania may seem unusual 57 (Stebnicka 2001) but not unheard of in other groups distributed in the southern hemisphere. The 58 59 60 Austral Entomology Page 26 of 81

26 1 2 3 purported case of longdistance transoceanic dispersal of the monotypic plant Myosotidium 4 5 hortensium , with a sister group occurring in the Mediterranean (with divergences between 3.64 6 7 and 22.45 Ma), may seem like a relictual quirk, but relationships among all the species that are 8 9 not shared with the mainland require intensive phylogenetic study. 10 11 12 13 14 The Deep South: the Subantarctic Islands 15 16 17 18 For Review Only 19 One of the best taxonomically documented group of the offshore islands are, surprisingly, the 20 21 Subantarctic Islands located on the Campbell Plateau, a large microcontinental fragment that 22 23 may have been a separate geological entity from the remaining northern portions of the mainland 24 25 New Zealand (Michaux & Leschen 2005). The New Zealand Subantarctic islands have attracted 26 the attention of biologists for well over a century, due to their isolation and importance for sea 27 28 life. There are five islands or archipelagos (Campbell, Auckland, Snares, Bounty, and Antipodes 29 30 Islands); this excludes Macquarie Island, which is located on the Macquarie Ridge which, though 31 32 its fauna is related to those of the Campbell Plateau, has few endemic species (Greenslade 1990). 33 The islands are scattered along the plateau, and range in size from the 135ha Bounty Islands to 34 35 the 61,120ha Auckland Islands, with all but the Bounty and Snares Islands being volcanic 36 37 remnants. Resting on Palaeozoic rocks, the oldest volcanic extrusions are 12–25 my (Adams 38 39 1981). Many of the islands are windswept, with Auckland Islands and Snares supporting forests 40 41 of Metrosideros and Olearia , respectively. There are up to 80% levels of endemism for some 42 groups of terrestrial invertebrates. While major surveys occurred in the 1960s, only a few have 43 44 been sporadically carried out since (see Gressit 1964; Gressit & Wise 1971). Some groups 45 46 require additional work (soildwelling insects, especially staphylinid beetles); and the fauna of 47 48 the Snares Islands has not been fully and formally documented. 49 50 Despite the thorough faunistic and taxonomic works of the 1960s and 1970s, 51 52 biogeography and evolution of the Subantarctic fauna, however, is surprisingly limited. Most 53 54 studies have argued that the Subantarctic faunas share relatives with mainland New Zealand, and 55 56 this includes taxa found also on the Chatham Islands (Kuschel 1971; Craw 1988; Michaux & 57 Leschen 2005; Ward et al . 2004; McGaughran et al . 2006; 2010; Leschen et al . 2011). Distant 58 59 60 Page 27 of 81 Austral Entomology

27 1 2 3 connections include taxa that have relatives in the northern Pacific ( Baeostethus chiltoni Broun; 4 5 Leschen et al . 2002; Ahn et al . 2010), but Gondwanan relationships predominate (Liebherr et al . 6 7 2011) with connections to the Patagonian regions of South America (Kuschel 1971) and to 8 9 Tasmania (Priasilphidae; Leschen & Michaux 2005). The internal biogeographic relationships of 10 11 the island invertebrates are poorly known. A phylogeny (Leschen et al . 2011) of the 12 Subantarctic beetle genus, Pseudhelops Guérin Méneville (Tenebrionidae), shows a lineage that 13 14 has colonised by western drifting, from the main group in the west to the Bounty Platform, which 15 16 is composed of relatively younger islands (Adams 1981).The evolutionary history among the 17 18 Campbell Plateau insectsFor remain poorlyReview studied, however, Only and much work is required to fully 19 appreciate the relationships of the insect fauna to other regions of the world, the relationships 20 21 among the islands, and especially their natural history. 22 23 24 25 26 GLACIAL REFUGIA IN NEW ZEALAND 27 28 29 History of glaciation in New Zealand 30 31 32 33 New Zealand’s history of repeated glaciations throughout the latePliocene and Pleistocene has 34 been particularly well characterised (e.g. Mercer 1983; Suggate 1990; Vandergoes & Fitzsimons 35 36 2003; Suggate & Almond 2005). Onset of cooling occurred in the Late Pliocene and intensified 37 38 throughout the Pleistocene, often rendering vast areas of the landmass as tundra. Repeating 39 40 cycles of glacial cooling and interglacial warm periods occurred during this period, driving 41 42 expansion and retreat of glaciers and significant sea level fluctuation (Suggate 1990). Indeed, the 43 Last Glacial Maximum (LGM), between 34 and 18 kya, saw extensive glaciation spanning 44 45 almost the entire Southern Alps, in concert with an approximately 120m reduction in sea level 46 47 (Newnham et al. 2012). These dramatic changes to New Zealand’s climate clearly drove 48 49 similarly marked transformations in terrestrial habitats across the landmass. Fossil data suggest 50 quite clearly that widespread, contiguous wet forests were fragmented by drier, open grassland 51 52 and tussock habitats and falls in sea levels exposed new land for colonisation, while 53 54 simultaneously converting previously coastal habitats into more inland zones (e.g. McGlone 55 56 1985; Newnham et al. 2012). It could be reasonably expected that many freshwater habitats 57 58 would have been assimilated by expanding glaciers or dried out completely and perhaps that 59 60 Austral Entomology Page 28 of 81

28 1 2 3 previously drier regions may have experienced increases in rainfall due to local changes in 4 5 climate. Although PlioPleistocene glaciation was extensive and changes to floral communities 6 7 significant, it is generally held that the landmass possessed a mosaic of vegetation types, rather 8 9 than complete conversion of broad regions and extirpation of core species (McGlone et al. 2010). 10 11 Nevertheless, this dramatic and dynamic fluctuation and conversion of habitat types also 12 influenced distributions and population connectivity in many resident insect groups. This section 13 14 details regions of the New Zealand landmass supported as refugia for insect taxa during past 15 16 glacial periods. 17 18 For Review Only 19 What makes a refugium? 20 21 22 23 Glacial refugia are recognised globally as land areas that represent formerly widespread habitats 24 25 that retracted in range during historical periods of glacial advance (Ferris et al . 1999). Typically, 26 though not the rule, terrestrial habitats that retracted into small refugia during glacial periods 27 28 comprised wet, closed canopy forest taxa that were driven locally extinct elsewhere by the 29 30 cooling and aridification that characterises glaciation. As a general result, refugial areas often 31 32 retained greater diversity and endemism than glaciated regions (Hewitt 2000; Byrne 2008); 33 34 although zones of overlap between refugia where divergent lineages recontact can harbour higher 35 diversity than individual refugia alone (Byrne 2008; Provan & Bennett 2008; Bryant & Fuller 36 37 2014). This trend holds for New Zealand, where a previously widespread closed forest flora was 38 39 fragmented into a mosaic of small isolated refugial areas. Despite New Zealand glaciation 40 41 occurring in mountainous areas instead of vast lowland ice shields (Suggate 1990), such refugia 42 were often (although not exclusively) located in upland areas or in valleys and gorges resistant to 43 44 the aridification of lowland areas. The dramatic transitions in floral communities across New 45 46 Zealand during glacial periods are welldocumented in the pollen record, particularly for the late 47 48 Pleistocene. In some areas this appears to have involved turnover in species composition as 49 50 changing local climates drove conversion from one habitat type to another (e.g. Nothofagus 51 dominated forest to open grassland), whereas in other areas floral communities appear to have 52 53 been extirpated by glacial advance (Wardle 1963; McGlone 1985). In contrast, subfossil beetle 54 55 remains from across New Zealand appear to imply that habitat conversion was not as dramatic as 56 57 inferred from the pollen record, with many Pleistoceneage fossil deposits suggesting persistence 58 59 60 Page 29 of 81 Austral Entomology

29 1 2 3 of a mosaic of small forest patches (reviewed by Marra 2008; Marra & Thackray 2010). A 4 5 further confounding factor for interpreting glacial age signals in New Zealand is that reductions 6 7 in sea level may have provided opportunities for lowland taxa to establish refugia in areas 8 9 currently inundated. 10 11 12 North Island – a line in the land 13 14 15 16 The extensive pollen record for the New Zealand North Island suggests the major centres of 17 18 floristic richness duringFor PlioPleistocene Review glacial periods wereOnly in Northland, Waikato (especially 19 Coromandel Peninsula), and the very southeastern tip of the island around the Tararua Ranges 20 21 (Wardle 1963, 1988). Indeed, in a recent metaanalysis of fossil data, Northland and Waikato are 22 23 the only regions in New Zealand supported as retaining significant tall podocarp forest during the 24 25 LGM (McGlone et al. 2010). Moreover, much of the southern half of the North Island below 26 around 39°S possesses markedly lower diversity and endemism than the northern half. This 27 28 dividing line has been termed the ‘Taupo Line’ after the settlement and lake which it intersects in 29 30 the central North Island. It is a zone of much contention among New Zealand biogeographers, 31 32 and no consensus for its development exists. Hypotheses include longterm marine transgression 33 34 (Suggate et al. 1978; Bunce et al . 2009), widespread tectonic activity (Pulford & Stern 2004) 35 and/or volcanism (Alloway et al . 2007). A glacial explanation has also been provided for the 36 37 Taupo Line that invokes retraction of taxa from southern areas to northern refugia (Wardle 38 39 1963). Whatever the case, the pattern suggests there has been little colonisation of much of 40 41 southern North Island since the LGM, implying that even if the drivers of the boundary are old, it 42 is being maintained by more modern processes. 43 44 Insects of the North Island largely reflect the floral data in showing higher levels of 45 46 diversity (both intra and interspecific) in northern zones, along with a small area in the 47 48 southeast, compared with the low diversity zone below the Taupo line. Ecological niche 49 50 modelling (ENM) of the generalist parthenogenetic stick insect, Argosarchus horridus (White), 51 suggests this taxon survived Pleistocene glaciations in refugia along coastal North Island, 52 53 including the southeast, and northeast North Island (Buckley et al . 2009). Mitochondrial DNA 54 55 data showed one common widespread clade distributed across South Island and southern North 56 57 Island, and several rarer clades mostly restricted to northern North Island. Furthermore, most 58 59 60 Austral Entomology Page 30 of 81

30 1 2 3 South Island populations were femaleonly, while North Island populations had almost equal sex 4 5 ratios – interpreted as resulting from southward colonisation of habitat during interglacials and a 6 7 selective advantage for parthenogenesis at the invasion front. This pattern is virtually mirrored in 8 9 another generalist parthenogenetic stick insect, Clitarchus hookeri (White), for which ENMs and 10 11 mtDNA data also suggest refugia in northern North Island and eastern South Island (Buckley et 12 al. 2010). Like A. horridus , northern populations of C. hookeri are sexual and genetically 13 14 diverse, whereas southern populations are parthenogenetic and of low diversity (Buckley et al. 15 16 2010; MorganRichards et al . 2010). The forest edge cicada species, Kikihia subalpina (Hudson), 17 18 showed a split across theFor Cook Strait Review for mitochondrial DNAOnly (Marshall et al. 2009), with North 19 Island populations most diverse in the southeast and a single lineage distributed north of the 20 21 Taupo Line. Refugial areas for two further taxa, the fungus beetle species Epistranus lawsoni 22 23 and Pristoderus bakewelli ¸ were supported by ENMs for northern and southeastern North Island, 24 25 including significant land areas in the north that are now below sea level (Marske et al. 2011). 26 Although little is known of refugial areas for North Island aquatic taxa during glacial periods, 27 28 some population genetic studies suggest their persistence in central and northern North Island 29 30 refugia as a driver of population divergence (Smith et al. 2006a,b). Taken together, insect data 31 32 for North Island match closely the interpretation provided for the pollen record and observed in 33 34 other terrestrial fauna: the northern portion of the island above the Taupo Line and the 35 southeastern tip were clearly major refugial areas for many taxa during Pleistocene glaciation. 36 37 38 39 South Island – montane mosaic 40 41 42 The topographically diverse South Island experienced major periods of glaciation several times 43 44 since the Pliocene. Glacial advance was mostly confined to the Southern Alps mountain range; 45 46 however, significant glaciation of lowland areas, particularly to the west, also occurred, while 47 48 much of central and eastern South Island supported mostly open grassland and tussock 49 50 (McGlone et al. 2010). Indeed, some fossil evidence suggests only northern South Island 51 (NelsonMarlboroughKaikoura), Westland, and a few small isolated areas of Fiordland 52 53 supported any significant tall forest during the LGM (McGlone et al. 2010). Nevertheless, other 54 55 evidence suggests Otago may have retained refugial areas during glaciations (Wardle 1963; 56 57 Burrows 1965), as may have areas of Canterbury (Marra et al. 2006). Contemporary patterns of 58 59 60 Page 31 of 81 Austral Entomology

31 1 2 3 floral communities have also informed inferences of the effect of PlioPleistocene glacial events 4 5 and, like the Taupo Line in North Island, areas of biotic disjunction has been recorded 6 7 throughout New Zealand, especially in the South Island. Much of the central South Island has 8 9 lower levels of species richness and endemism than adjacent northern and southern areas (Willet 10 11 1950; Wardle 1963). This “Biotic Gap” is likely the result of several ecological and historical 12 processes (e.g. Heads 1998; Trewick & Wallis 2001; Haase et al . 2006). Of particular note is the 13 14 “Westland beech gap”, a 150 km long region on the West Coast of South Island that lacks beech 15 16 trees (e.g., Cockayne 1926; Wardle 1963; McGlone et al . 1996) and a number of insect taxa 17 18 (e.g., Leschen et al . 2008).For Review Only 19 Refugial areas inferred for South Island insects based on molecular phylogeographic 20 21 studies largely follow predictions made from fossil data. The widely distributed fungus beetles, 22 23 Brachynopus scutellaris (Redtenbacher) and Hisparonia hystrix (Sharp), show high intraspecific 24 25 genetic diversity with divergent lineages distributed across the welldemarcated Westland Beech 26 Gap, but with substructuring of lineages somewhat correlated with geography (Leschen et al 27 28 2008). Patterns in both species are inferred to have resulted from a complex history of population 29 30 fragmentation and reexpansion associated with glacial cycling. The stick insect Niveaphasma 31 32 annulata (Hutton) comprises three Pleistoceneage lineages restricted to putative refugial areas 33 34 in Canterbury, Otago, and FiordlandSouthland (O’Neill et al. 2009). Distributions of several 35 species of South Island tiger beetles ( Neocicindela Rivalier) appear centred around historical 36 37 refugial areas, especially NelsonMarlborough, Canterbury, Otago, and the Southern Alps (Pons 38 39 et al . 2011). Many Neocicindela species that are distributed across the Biotic Gap also show 40 41 deep divergence between northern and southern populations and genetic signals suggest the 42 Biotic Gap was recolonised by these taxa during interglacial periods. A study of the forest fungus 43 44 beetle, Argyrtodes labralis (Broun), that incorporated ENMs with traditional molecular 45 46 phylogeography suggested Pleistocene refugia for this taxon would have been located in northern 47 48 South Island and the west coast, along with a very restricted zone in Fiordland (Marske et al. 49 50 2009). The molecular data supported this in showing several highly localised lineages in 51 Kaikoura, Nelson, and Marlborough, with some others more widespread in OtagoSouthland and 52 53 the west coast. Southern regions evidently were not completely extirpated during glaciations, 54 55 with populations surviving in small zones of suitable habitat, and that west coast refugia were the 56 57 source populations for eastern and southeastern populations, with very little movement out of 58 59 60 Austral Entomology Page 32 of 81

32 1 2 3 Nelson refugia (Marske et al. 2009). The cicada K. subalpina , mentioned above, showed 4 5 significant substructuring among South Island populations, with greatest diversity in the northern 6 7 refugial regions and little diversity in southern areas (Marshall et al. 2009). Both K. subalpina 8 9 and A. labralis also inhabit Stewart Island and the surveys described above suggest Stewart 10 11 Island is likely to have been colonised only since the LGM by populations in southern South 12 Island (Marshall et al. 2009; Marske et al. 2009). 13 14 Some freshwater insects in South Island also show distinct histories of isolation in refugia 15 16 during glacial cycles. All six codistributed species of stonefly assessed by McCulloch et al. 17 18 (2010) were concordantFor in showing Review deep genetic divergences Only across the Biotic Gap, with few 19 genetic lineages distributed across the Gap, and centres of diversity correlated with putative 20 21 refugial areas. Some of the studied species currently inhabit the Biotic Gap and the authors 22 23 suggest dispersal ability (as determined in this case by aptery vs macroptery) as the crucial factor 24 25 in facilitating recolonisation of glaciated areas (McCulloch et al. 2010). Populations of the non 26 biting midge, Naonella forsythi Boothroyd, in the Upper Buller catchment appear to have 27 28 experienced significant subdivision, divergence and subsequent reexpansion (Krosch et al. 29 30 2012), possibly due to glaciation in the region (Suggate 1990). The waterboatman species, 31 32 Sigara potamius Young, also exhibits marked genetic divergence in northern South Island, with 33 34 some lineages restricted to locations on the west coast and others sympatric in Canterbury 35 (Buckley & Young 2008). Diversification in this species was inferred to have initiated in the 36 37 Pliocene and it is likely that refugia in northern South Island were important for persistence 38 39 during the Pleistocene. 40 41 In contrast to the evidence above, some taxa are supported as having survived through the 42 Pleistocene glaciations in refugia within the Biotic Gap. The cicada, Maoricicada campbelli 43 44 (Fleming), is suggested to have persisted in the Biotic Gap and diversified in Pleistocene refugia 45 46 in southern North Island, Marlborough, the Southern Alps, Waitaki, and Central Otago (Buckley 47 48 et al. 2001a; Hill et al. 2009). 49 50 Confounding interpretations of glacialage population impacts is the finding that even 51 ecologically similar taxa may respond differently to glaciation, having inhabited different 52 53 refugial areas, used different recolonisation paths and diversified at different tempos (see Marske 54 55 et al. 2012). Furthermore, many taxa that may have inhabited refugia in the Pleistocene may 56 57 have lost any genetic signal over the intervening time due to other processes (both intrinsic and 58 59 60 Page 33 of 81 Austral Entomology

33 1 2 3 extrinsic) and many different geophysical processes may produce the same phylogeographical 4 5 patterns (Trewick et al. 2011). Despite this, and the fact that many taxa apparently were not at all 6 7 affected by Pleistocene glaciations (Trewick et al. 2011) and several insects are endemic to the 8 9 Westland Beech Gap (e.g. Leschen & Michaux 2005; Seago & Leschen 2011), there remains a 10 11 clear signal in the evolution of numerous New Zealand insect species of significant impact on 12 populations due to glacial advance. Questions still remain concerning the influence of glaciations 13 14 in many insect groups and ongoing research is needed to elucidate these effects. Understanding 15 16 the impact of past climatic fluctuations will facilitate effective prediction and management of 17 18 future change. For Review Only 19 20 21 22 CONSERVATION OF NEW ZEALAND INSECTS 23 24 25 26 New Zealand insects under threat 27 28 29 The remarkable diversity, endemism and evolutionary histories of New Zealand’s insect fauna 30 31 pose many conservation problems in a changing humaninfluenced landscape. New Zealand is 32 33 considered a biodiversity hotspot for conservation and possesses a community of species unique 34 in the world (Myers et al. 2000). The oftendevastating trident of exotic species, habitat 35 36 loss/modification (particularly in the lowlands), and climate change represent increasing 37 38 challenges for the survival of many insect species in New Zealand. Current estimates suggest 39 40 that of 3,838 terrestrial invertebrate species assessed (26% of the New Zealand total), 193 were 41 42 considered ‘Threatened’ (including 106 that were ‘Critical’), and a further 1,055 were ‘At Risk’ 43 (Stringer & Hitchmough 2012). Moreover, 912 species ‘At Risk’ of extinction were considered 44 45 ‘Naturally Uncommon’, an important contributor to their conservation status and meaning that 46 47 otherwise minor impacts could have a dramatic influence on these taxa. Around 114 species of 48 49 New Zealand Lepidoptera are considered of conservation significance (Patrick & Dugdale 2000), 50 and many of the large, ground dwelling weta are known to be vulnerable to predation (Gibbs 51 52 1998). Already seven insect species are believed recently extinct and others exist now only on 53 54 smaller islands (Samways 1993; Gibbs 2009). Furthermore, many more species remain 55 56 undescribed and with unknown conservation status, and thus the true extent of the threat may be 57 58 greater than currently thought (McGuiness 2001; Stringer & Hitchmough 2012).All is not lost, 59 60 Austral Entomology Page 34 of 81

34 1 2 3 however; this section will explore some important conservation issues for terrestrial and aquatic 4 5 taxa, threats, solutions, success stories and the future prospects for New Zealand’s insects. 6 7 8 9 Standing on solid ground 10 11 12 Exotic species have had and will continue to have an enormous impact on native terrestrial insect 13 14 populations across New Zealand. Several wellknown mammalian invaders directly predate on 15 16 insects, including rodents, possums, mustelids, pigs, and hedgehogs (McGuiness 2001; Gibbs 17 18 2009). Several weevil Forand beetle species,Review including the largestOnly known ulodid beetle 19 (Waitomophylax worthyi [Leschen & Rhode]), are thought to have been driven to extinction from 20 21 the mainland or altogether due to rat predation (Kuschel & Worthy 1996; Leschen & Rhode 22 23 2002; Gibbs 2009; Leschen et al. 2012). Throughout the New Zealand archipelago rats have also 24 25 been a significant causative agent in the decline of many weta species, especially those large taxa 26 that spend much time on the ground and have inadequate defensive strategies (Gibbs 1998, 27 28 2009). So effective have rats and other mammalian predators been in reducing mainland 29 30 populations of such insects, that a number of these insects have been deliberately translocated to 31 32 mammalfree islands to prevent further decline (Watts et al. 2008). Among the translocated 33 34 species is the Mercury Islands tusked weta, Motuweta isolata Johns; the entire extant population 35 of which is descended from three individuals. Since then, individuals from a captive breeding 36 37 programme were released successfully onto two islands in the Mercury group (Double and Red 38 39 Mercury Islands) to reduce the potential for accidental extinction on the only island they were 40 41 known from originally (Middle Island) and from which they have not been recorded for several 42 years (since January 2001) (Stringer & Chappell 2008). Of the other mammal and avian invaders 43 44 that predate on native insects, little appears to be known of the extent of their impact on 45 46 populations and it could be assumed that it may be minimal compared with the impact of rats. 47 48 Some introduced insects also predate on native taxa, the most well known are the social 49 50 wasps of the genus Vespula Thomson. Two species of this genus, V. vulgaris (Linnaeus) and V. 51 germanica (Fabricius), have established in New Zealand and both are known to predate on native 52 53 Diptera, Lepidoptera, and Aranae (Beggs & Rees 1999). Wasp populations have been shown to 54 55 reach incredible sizes in native forest and can cause dramatic population declines and sometimes 56 57 local extinctions of native spiders (Toft & Rees 1998) and butterflies (Beggs & Rees 1999). 58 59 60 Page 35 of 81 Austral Entomology

35 1 2 3 Wasps also utilise the abundant carbohydrate resources from native scale insect exudate or 4 5 honeydew (up to 70% of standing honeydew production). This contributes to competition with 6 7 other honeydew users, including birds and other insects. Vespula wasps can dramatically re 8 9 structure forest communities during peak wasp season (JanuaryApril) as prey species that 10 11 normally occur are extirpated. Taken together, social wasps represent one of the chief 12 conservation issues concerning exotic invaders for native New Zealand insects. 13 14 Some other introduced invertebrates influence native insects through competition, 15 16 parasitism and predation. For example, the Southern African mantis ( Miomantis caffra Saussure) 17 18 has been spreading fromFor Auckland, Review where it was recorded Only first in 1978, and displacing the native 19 mantis, Orthodera novaezealandiae (Colenso) (see Brockerhoff et al. 2010). A few predacious 20 21 beetles, parasitoid social insects, and ants are established in New Zealand but are rare in native 22 23 forest and their impact on native insects is thought to be minimal. Likewise, some biocontrol 24 25 agents have been recorded to exhibit nonspecific attack (parasitoid wasps supposed to attack 26 crop and pasture pests also attack native Lepidoptera, weevils); however, their impact on native 27 28 taxa may yet be minimal – in contrast to many other areas of the world (Brockerhoff et al. 2010). 29 30 Surprisingly little is known of the impact of invasive plants on native insect populations; 31 32 although there is some evidence that the invasive herb Tradescantia fluminensis reduces 33 34 abundance of epigaeic invertebrates (Standish 2004), but has no impact on beetles or fungus 35 gnats (these instead correlated with vascular plant richness and diversity regardless of presence 36 37 of exotic flora – Crisp et al. 1998; Toft et al. 2001). 38 39 Aside from introduced species, perhaps the greatest threat to native terrestrial insect 40 41 populations and communities remains habitat loss and modification through human impact, with 42 some groups confined to small areas (Cromwell Chafer, Prodontria lewisi Broun, Watt 1979) or 43 44 forest fragments, like Radar Bush in Northland (Buckley & Bradler 2010). This can be through 45 46 either wholesale change to a particular parcel of land (e.g. through land clearing) or more minor 47 48 changes that have ripple effects through the ecosystem (e.g. erosion, trampling, and grazing of 49 50 native vegetation by livestock). Formerly, subsidies were provided by the New Zealand 51 government and access to cheap loans given to individuals to clear forest to expand agricultural 52 53 enterprises and for the woodchipping industry (Howarth & Ramsay 1991). Thankfully, the 54 55 removal of these incentives has significantly slowed the decline and loss of habitat. Nevertheless, 56 57 threats from these processes remain and native habitats continue to be disturbed by human 58 59 60 Austral Entomology Page 36 of 81

36 1 2 3 practices (be they agriculture, forestry, mining or urban development). It remains unclear 4 5 whether New Zealand’s insect fauna is suitably protected to cope with these ongoing threats; 6 7 however, recent work suggests the endangered ground beetle, Holcaspis brevicula Butcher, 8 9 whose native range has been fragmented by human practices, has expanded out into forestry 10 11 plantations that inadvertently provide suitable habitat for it and other beetles (Brockerhoff et al. 12 2005; Pawson et al . 2008). Clearly some of New Zealand’s insects can adapt to ongoing human 13 14 influence, but we expect this phenomenon to be the exception. 15 16 17 18 The life aquatic For Review Only 19 20 21 The conservation status of many New Zealand freshwater insects is poorly known, and despite a 22 23 review of this theme in the 1990s (Collier 1993), little new information on this topic has come to 24 25 light in the years since. Collier (1993) recognised 154 species of insects inhabiting surface 26 waters that possess restricted distributions, as well as 20 species from underground systems and 27 28 36 isolated on offshore islands. Localised distributions clearly put taxa at risk of extinction due 29 30 to habitat change and more data are needed to appropriately assess risks to these taxa. Of those 31 32 threats to freshwater taxa that have been established, it is abundantly clear that habitat loss and 33 34 modification is the greatest cause for concern. Loss and modification of freshwater ecosystems 35 can take many forms and include: degradation and loss of riparian zones, channel environments 36 37 and catchment properties, changes to flow regimes, declines in water quality (increased sediment 38 39 load, decreased pH, increased nutrient load, altered thermal regimes), and increases in pollution 40 41 (Collier 1993). Often compounding effects of such impacts is the dendritic nature of river 42 networks that means upstream impacts are cumulative on downstream sections. There are several 43 44 driving factors behind the modification of aquatic environments, not least of which are 45 46 urbanisation, mining, forestry, and agriculture. Moreover, grazing of livestock has also been 47 48 shown to have a dramatic effect on bank damage, riparian degradation, and associated increased 49 50 sediment load – all of which have a marked effect on insect communities (Quinn et al. 1992). 51 New Zealand’s freshwater taxa have also been dramatically affected by exotic invaders, 52 53 especially fish (most notably brown trout, Salmo trutta L.), snails, and macrophytes. The 54 55 introduction in the mid–late 1800s of brown trout and other salmonids to New Zealand 56 57 waterways for the angling community is thought to have increased predation pressure on some 58 59 60 Page 37 of 81 Austral Entomology

37 1 2 3 larger native aquatic invertebrates, including mayfly and stonefly nymphs (Collier 1993). The 4 5 invasive snail, Physa acuta (Draparnaud), has displaced a native species in many areas 6 7 (Winterbourn 1973). Exotic aquatic flora cause changes to water quality, contribute to habitat 8 9 degradation, and alter flow regimes by clogging waterways. Invasive macrophytes have also 10 11 been shown to significantly alter species composition in the littoral zone of lakes and increase 12 the total density, richness, and diversity of insect inhabitants (Kelly & Hawes 2005). In contrast, 13 14 clogging of waterways by some aquatic weeds can sometimes support an increase in abundance 15 16 in native snails, Trichoptera and Diptera, possibly by providing an additional food resource, but 17 18 also perhaps because theFor weeds excludeReview predators and provide Only secluded areas for larval 19 development. 20 21 22 23 Future prospects and the spectre of changing climate 24 25 26 New Zealand, paradoxically, is wellplaced to deal successfully with conservation issues 27 28 concerning its insect fauna (New & Samways 2013). The public, over many years, has embraced 29 30 conservation principles and become enamoured with many of New Zealand’s iconic insect 31 32 species such as glowworms and weta (Cranston 2010). Furthermore, the Department of 33 34 Conservation (DOC) has been instrumental in managing many of the risks discussed above. For 35 example, although a significant portion of New Zealand’s native habitats are privately owned 36 37 and of uncertain conservation status, protected areas now constitute around 30% of the landmass, 38 39 including a growing number of ‘mainland islands’, managed by DOC (Saunders & Norton 40 41 2001), local government, trusts, and various citizens groups. This is a remarkable achievement 42 and serves to highlight the importance placed on maintaining significant blocks of continuous 43 44 habitat. Likewise, urban reserves (smaller patches of native forest in an urbanised landscape) 45 46 have been shown to be a significant potential refuge for insect diversity in an increasingly 47 48 modified landscape, thus highlighting the importance of green areas in the urban mosaic (Watts 49 50 & Lariviere 2004). Of course, passive protection is often insufficient to combat the decline of 51 native species (Clout 2001) and even within protected areas exotic species roam freely. 52 53 Mammal invaders are trapped and baited throughout the mainland to control numbers and 54 55 eradication programmes have been successful on many offshore islands and within the mainland 56 57 islands (Saunders & Norton 2001; Clout & Russell 2006). For example, removal of rats and 58 59 60 Austral Entomology Page 38 of 81

38 1 2 3 weka from the Chetwode Islands in Pelorus Sound resulted in increases in the proportion of 4 5 active adult weta in a given habitat and the proportion of adults in the population, along with 6 7 changes to population dynamics and habitat use associated with lowered predation (Rufaut & 8 9 Gibbs 2003). Several methods of control have been implemented for Vespula wasps, including 10 11 biocontrol using parasitoid wasps (Beggs & Rees 1999) and poison sprays (Beggs et al. 1998, 12 Toft & Rees 1998); however, none has succeeded on broad spatial scales (Beggs 2001). Control 13 14 methods for many invasive plants currently are often longterm and laborious, involving 15 16 mechanical removal, spraying of often potent herbicides, monitoring, and sometimes grazing 17 18 with livestock to preventFor reemergence Review (Ogle et al. 2000; Only Hofstra et al 2001). In contrast, 19 biocontrol measures for invasive plants are considered to be quite effective in New Zealand 20 21 (Fowler et al. 2000). 22 23 Arguably the most pressing concern for New Zealand’s insect fauna, and particularly the 24 25 alpine taxa, is the influence of climate change. Already, evidence for the influence of changing 26 climate is abundant in New Zealand, with dramatic reductions in many glaciers (Chinn 1996) 27 28 and rising timberlines (Wardle & Coleman 1992). Unfortunately, there has been little work 29 30 conducted to identify directly the likely impacts on native insects from future climate change; 31 32 however, lessons can be learned from historical patterns outlined in previous sections. If future 33 34 climate change drives higher temperatures and reduced rainfall across New Zealand as appears to 35 be the case (Plummer et al. 1999), it is anticipated that susceptible habitats will respond as in the 36 37 past. That is: closed, wet forest will retract into isolated refugia; dry grassland will expand across 38 39 the lowlands; coldadapted taxa will be limited to mountaintops; and, in the extreme case, taxa 40 41 that cannot shift distribution will be driven to extinction (Wardle & Coleman 1992). Moreover, 42 climate change may also indirectly impact on freshwater habitats via increased temperatures and 43 44 more frequent droughts that increase sediment loads and reduce flows. It has been suggested that, 45 46 depending on the degree of temperature rise, several hundred alpine vascular plant taxa could be 47 48 lost, along with a substantial proportion of existing alpine ‘islands’ (Halloy & Mark 2003). 49 50 Taken together, this clearly presents a significant risk not just to the alpine insect fauna but to 51 that of New Zealand as a whole. There exists a growing need for fundamental research into the 52 53 potential impacts of climate change on New Zealand insects and whether there are any suitable 54 55 mitigation strategies that could be put in place. The incredible endemism of New Zealand’s 56 57 58 59 60 Page 39 of 81 Austral Entomology

39 1 2 3 insect fauna is worth protecting and concerted effort should be put into ensuring their survival 4 5 into the future. 6 7 8 9 10 PROSPECTUS 11 12 13 14 15 We have reviewed some of the major historical processes that have shaped the diversity and 16 17 special features of the New Zealand insect fauna. While great strides have been made in 18 understanding the origins,For diversification, Review and adaptive evolutionOnly of the insect fauna over recent 19 20 years, there are still significant gaps in our knowledge. Importantly, these gaps impede our 21 22 ability to manage and protect the insect fauna for future generations. However, there are also 23 24 great opportunities, not only for increasing knowledge and understanding of individual lineages, 25 26 but also for using New Zealand insects as model systems to test more generally outstanding 27 hypotheses from the field of evolutionary biology. 28 29 30 One of the most critical knowledge gaps is taxonomic: the insect fauna remains too 31 st 32 poorly known (Leschen et al . 2009; Lester et al . 2014). Even in the 21 Century, large insects 33 from the usually better known terrestrial macrofauna, such as large orthopteroids (e.g. Jewell 34 35 2007; Buckley & Bradler 2010; Taylor Smith et al . 2013), are being discovered and described. 36 37 The taxonomic knowledge gaps in the hyperdiverse microfauna are orders of magnitude greater. 38 39 Furthermore, there are also significant gaps in our understanding of the distributions of species 40 41 and patterns of species richness at local and national scales. This can only be remedied with 42 specialised collecting techniques, which are required for most insect groups (e.g. Seldon & 43 44 Beggs 2010), systematics research, and dramatically increasing the degree of curation and data 45 46 accessibility in collections (e.g. Ward 2012). Targeted surveys of poorly known areas, including 47 48 the alpine zone, are required urgently and our own collecting on offshore islands, often 49 considered “well collected”, frequently yields new species and even genera (e.g. Théry & 50 51 Leschen 2013). Family group records that are new to New Zealand are made regularly (e.g. 52 53 Grebennikov & Newton 2008; Ruta et al . 2011). Only by building the taxonomic framework 54 55 along with comprehensive information on geographic distributions will we be in a position to 56 57 construct and test detailed hypotheses on the evolution of large elements of the insect fauna. 58 59 60 Austral Entomology Page 40 of 81

40 1 2 3 There are essentially endless opportunities for advancing taxonomic knowledge on the New 4 5 Zealand insects. 6 7 8 A robust phylogenetic framework is a prerequisite for any evolutionary study (e.g. 9 10 Felsenstein 1985), and numerous phylogenies have emerged from studies on New Zealand 11 insects that are discussed and cited in this review. However, there is a clear bias in these studies 12 13 towards the insect macrofauna, such as weta, stick insects, cicadas, and the larger beetles. 14 15 Clearly neglected are hyperdiverse groups, especially those associated with dead wood, leaf 16 17 litter, and soil microhabitats, including Oecophoridae moths, Microgastrinae wasps, a number of 18 Diptera families, and manyFor Coleoptera Review groups, especially Only Staphylinidae, Curculionidae, 19 20 Tenebrionidae, and Zopheridae (see Watt 1975; Klimaszewski et al . 1996; Lord & Leschen 21 22 2014). Even among less diverse invertebrate groups (e.g. Leschen 2006; Boyer & Giribet 2010) 23 24 from the soil, leaf litter, and dead wood habitat, only a few groups have been examined 25 26 phylogenetically with absolute or relative divergence time estimation and comprehensive 27 sampling of relevant lineages from other land masses to determine their biogeographic origins, 28 29 including Cyphophthalmi mite harvestment (Boyer & Giribet 2009), Onychophora (Murienne et 30 31 al . 2014), and earthworms (Buckley et al . 2011). None of these groups are from the Insecta, 32 33 despite the hyperdiversity of insects from soil and saproxylic habitats in New Zealand (e.g. 34 Hoare 2005, 2010; Leschen 2006; Lord & Leschen 2014). We believe that such groups will 35 36 contain a greater number of archaic lineages relative to more arboreal, winged insects from the 37 38 macrofauna, many of which show more recent ancestry (e.g. Brown et al . 1999; Arensburger et 39 40 al . 2004). Given that the microfauna contain the bulk of New Zealand’s insect diversity, it is 41 42 premature to infer general patterns about the origin and evolution of the insect fauna, until well 43 supported phylogenetic reconstructions from these groups are obtained. 44 45 46 Extensive progress can also be made through exploiting the information held across the 47 48 existing, multiple phylogenies. Modelbased biogeographic methods exist that are able to draw 49 50 general patterns from simultaneous analysis of multiple phylogenies that include taxa distributed 51 across common areas (e.g. Sanmartín & Ronquist 2004; Ree & Sanmartín 2009). Comparative 52 53 biogeographic methods used by Sanmartín and Ronquist (2004) to infer different biogeographic 54 55 histories across sets of taxa from the Southern Hemisphere are exemplary. Metaanalyses 56 57 examining dozens of phylogenies for New Zealand insects and other organisms will allow 58 59 60 Page 41 of 81 Austral Entomology

41 1 2 3 detection of what organismal characteristics, such as taxonomic group, dispersal ability, and 4 5 ecological trophic level have most strongly influenced insect biogeographic history. 6 7 8 A major technical challenge for gaining greater insight into the ages of different insect 9 10 lineages in New Zealand is obtaining accurate divergence times. The problematic nature of this 11 has been well covered elsewhere, but includes problems with uncertainty and error in branch 12 13 length estimation (Buckley et al . 2001b), selection and testing of an appropriate relaxed clock 14 15 model (e.g. Baele et al . 2013), and the confounding effects of extinction (e.g. Waters & Craw 16 17 1996; Sharma & Wheeler 2013). However, it is widely acknowledged that the greatest potential 18 source of error in molecularFor dating Review studies is the reliability Only of calibrations (Parham et al . 2012). 19 20 Because the New Zealand fossil insect fauna is relatively poor, largely restricted to Quaternary 21 22 deposits (e.g. Marra 2008) but with a growing number of records from the late Oligocene and 23 24 early Miocene (Kaulfuss et al . 2010), there are few opportunities to internally calibrate 25 26 phylogenies. One potential solution to this problem is to integrate New Zealand and relevant 27 outgroup lineages into global phylogenies of higher level taxa that are often calibrated with a 28 29 larger number of fossils. This is because it is easier to assign fossils to a higher level taxon than a 30 31 lower level taxon such as a genus. This approach was taken by Perrie and Brownsey (2007) to 32 33 establish the ages of a range of New Zealand fern lineages. The growing number of large 34 phylogenomic data sets with multiple fossil calibrations offers an opportunity to achieve this for 35 36 New Zealand insect groups. A related approach in the absence of internal fossil taxa is to use an 37 38 estimated date for the age of a higher level taxon from a multicalibration phylogenomic study to 39 40 constrain the age of that lineage in a molecular dating study of New Zealand lineages. This 41 42 approach was used by Murienne et al . (2014) to date the age of lineages of Onychophora, 43 including those from New Zealand. The rapidly growing number of wellcalibrated, 44 45 phylogenomic studies offer great potential to gain increased knowledge of the absolute ages of 46 47 New Zealand lineages and then relate these ages to various ecological and historical processes. 48 49 50 A further promising avenue of research is to explore the characteristics of lineages that 51 have enabled them to speciate. This requires detailed understanding of the adaptations allowing 52 53 species to survive in their environments in addition to the characters that maintain species 54 55 integrity. Two study systems that have provided deep insights into basis of New Zealand insect 56 57 speciation are the cicada genus Kikihia Dugdale and the native tortricid moth genera Planotortrix 58 59 60 Austral Entomology Page 42 of 81

42 1 2 3 Dugdale and Ctenopseustis Meyrick. Like all other cicadas, Kikihia species have species specific 4 5 songs (e.g. Fleming & Dugdale 1984; Marshall et al . 2008). Molecular phylogenetic studies have 6 7 shown that these songs can evolve rapidly in isolated populations (Marshall et al . 2011). 8 9 Although song differences correlate well with genetic lineages, in many cases these do not 10 11 appear to have completely isolated species, with hybrid zones characterised by introgression of 12 mitochondrial DNA and intermediate song phenotypes being observed (Marshall et al . 2008, 13 14 2011). Similarly, species within the moth genera Planotortrix and Ctenopseustis can be defined 15 16 on the basis of unique pheromone blends (Newcomb & Gleeson 1998), yet the species are very 17 18 closely related (LanghoffFor et al . 2009). Review Genomic studies haveOnly shown that these differences are the 19 result of differential expression of desaturase genes involved in pheromone biosynthesis in 20 21 females (Albre et al . 2012). On the male side of this interaction, accelerated evolutionary rates of 22 23 odorant receptors in different lineages are likely responsible for tracking the evolution of the 24 25 female pheromone signal (Carraher et al . 2012). By examining the genetic basis of character 26 systems in other insect lineages, and placing this in a phylogenetic context, we will gain greater 27 28 insight into the processes that have generated the diversity of New Zealand insects. 29 30 31 Determining the most important organismal characteristics to have influenced 32 33 biogeography requires extensive knowledge on the biology and ecology of individual insect 34 species, natural history information that is often contained in systematics treatments and in 35 36 supplementary information in community ecology studies. Specific studies into the underlying 37 38 biology of various insect groups are underway, including diet and feeding ecology (e.g. 39 40 Kadowaki et al . 2011b; Larsen & Burns 2012; Wehi et al . 2013), cold tolerance physiology and 41 42 genetics (e.g. Sinclair et al . 1999; Dunning et al . 2014; Minards et al . 20014), symbiotic 43 relationships (e.g. Dhami et al . 2013), behavioural ecology (e.g. Kelly 2008; Painting & Holwell 44 45 2014), herbivory (e.g. Kelly et al . 1992), comparative phenology (e.g. Gibbs 2014) and 46 47 functional morphology (e.g. Painting et al . 2014; Yavorskaya et al. 2014). Detailed studies of 48 49 individual taxa lead to great insights, but contrasting traits among related species in a 50 phylogenetic framework will lead to the greatest advances in understanding the evolution of the 51 52 insect fauna. Such studies will enable us to answer questions concerning how often convergent 53 54 evolution occurs in species radiations, whether single character state (or key innovations) 55 56 transformations or combinations of characters have triggered adaptive radiations, and why some 57 58 lineages have radiated extensively compared with others that are depauperate (e.g. Losos 2010). 59 60 Page 43 of 81 Austral Entomology

43 1 2 3 The massive increase in the output of DNA sequencing technologies in the past 10 years 4 5 has been striking. This has significantly impacted all fields of biology, and evolutionary biology 6 7 is no exception (Ekblom & Galindo 2011; McCormack et al . 2013). High throughput DNA 8 9 sequencing allows a much greater number and range of markers to be applied to a given 10 11 organism through speciesspecific marker development (e.g. Abdelkrim et al . 2009) or more 12 direct methods such as ‘Genotyping by Sequencing’ (Elshire et al . 2011; Narum et al . 2013) and 13 14 sequence capture (e.g. Faircloth et al . 2012; Lemmon et al . 2012). The largescale data sets that 15 16 result from these technologies offer vastly improved estimates of phylogeny, divergence times, 17 18 genetic diversity, and Forgene flow withinReview and among species, Only and they are already being applied to 19 the New Zealand insect fauna (e.g. Hale et al . 2010; Dunning et al . 2013b, 2014). A particularly 20 21 promising research avenue that these methods open up is determining the genetic basis of 22 23 phenotypic traits and therefore improved understanding of their evolution (e.g. Storz & Wheat 24 25 2010; Dalziel et al . 2009). Methods such as RNA sequencing allow gene expression to be 26 measured in an organism under different treatment regimes, allowing inference of the genetic 27 28 basis to the response to specific environmental conditions (e.g. Dunning et al . 2014). Other 29 30 methods, such as RNA interference and in vivo expression in cell lines, allow the function of 31 32 specific genes to be directly determined (e.g. Albre et al . 2012), so the genetic basis of function 33 34 can be examined phylogenetically. The expansion of functional genomics to further elements of 35 the New Zealand insect fauna will facilitate significant advances in our understanding of the 36 37 evolution of adaptation and how insect species have evolved into unique New Zealand 38 39 environments. 40 41 42 In summary, no single method or avenue of research is a panacea for understanding the 43 origin and evolution of the New Zealand insect fauna, and a synthesis of systematics, ecological, 44 45 phenotypic and genomic research is required. The New Zealand fauna is an important part of the 46 47 global fauna, in age, diversity, and uniqueness. The wellcircumscribed geography and improved 48 49 understanding of geological history means New Zealand is wellsuited as a model ecosystem and 50 continues to attract a wide variety of scientists, locally and globally. New Zealand is also a 51 52 naturally inspiring place with a beautiful range of habitats across a rather small and accessible 53 54 land area, vast in its rich diversity and offering unlimited evolutionary questions. 55 56 57 58 59 60 Austral Entomology Page 44 of 81

44 1 2 3 ACKNOWLEDGEMENTS 4 5 6 We thank Pete Cranston, Mark Stevens and Anne Austin for comments on the text. TRB and 7 8 RABL were supported by core funding for Crown Research Institutes from the Ministry of 9 10 Business, Innovation and Employment's Science and Innovation Group. TRB also received 11 support from The University of Auckland, The Royal Society of New Zealand Marsden fund 12 13 (LCR0901) and Allan Wilson Centre. MNK was supported by a Postdoctoral Research 14 15 Fellowship awarded by the University of Queensland. We thank John Marris for unpublished 16 17 information on Three Kings beetles and Des Helmore for the illustrations. 18 For Review Only 19 20 21 REFERENCES 22 23 24 Abdelkrim J, Robertson BC, Stanton JAL & Gemmell NJ.. 2009. Fast, costeffective 25 26 development of speciesspecific microsatellite markers by genomic sequencing. 27 Biotechniques 46 , 185–192. 28 29 30 Albre J, Liénard MA, Sirey TM et al . 2012. Sex pheromone evolution is associated with 31 32 differential regulation of the same desaturase gene in two genera of leafroller moths. 33 PLoS Genet 8(1), e1002489. 34 35 36 Adams CJ. 1981. Migration of late Cenozoic volcanism in the South Island of New Zealand and 37 38 the Campbell Plateau. Nature 294 , 153–155. 39 40 41 Ahn K J, Jeon MJ & Branham MA. 2010. Phylogeny, biogeography and the stepwise 42 evolutionary colonization of intertidal habitat in the Liparocephalini based on 43 44 morphological and molecular characters (Coleoptera: Staphylinidae: 45 46 Aleocharinae). Cladistics 26 (4), 344–358. 47 48 Allegrucci G, Trewick SA, Fortunato A, Carchini G & Sbordoni V. 2010. Cave crickets and cave 49 50 weta (Orthoptera, Rhaphidophoridae) from the southern end of the World: a molecular 51 52 phylogeny test of biogeographical hypotheses. Journal of Orthoptera Research 19 , 121– 53 54 130. 55 56 57 58 59 60 Page 45 of 81 Austral Entomology

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75 1 2 3 Wehi PM, Raubenheimer D & MorganRichards M. 2013. Tolerance for nutrient imbalance in an 4 5 intermittently feeding herbivorous cricket, the Wellington tree weta. PLoS ONE 8(12), 6 7 e84641. 8 9 10 Wharton DA. 2011. Cold tolerance of New Zealand alpine insects. Journal of Insect Physiology 11 57 , 1090–1095. 12 13 14 Willet RW. 1950. The New Zealand Pleistocene snow line, climatic conditions, and suggested 15 16 biological effects. New Zealand Journal of Science and Technology 32 , 18–48. 17 18 Winterbourn MJ. 1973.For A guide toReview the freshwater Mollusca Only of New Zealand. Tuatara 20 , 141– 19 20 159. 21 22 23 Winterbourn MJ. 1980. The freshwater insects of Australasia and their affinities. 24 25 Palaeogeography, Palaeoclimatology, Palaeoecology 31 , 235–249. 26 27 Winterbourn MJ. 1997. New Zealand mountain stream communities: Stable yet disturbed? In: 28 29 Evolutionary Ecology of Freshwater Animals (eds B Streit, T Städler & C Lively). 30 31 Birkhäuser, Basel. 32 33 Winterbourn MJ. 2004. Association between a commensal chironomid and its mayfly host in 34 35 rivers of North Westland. New Zealand Natural Sciences 29 , 21–31. 36 37 38 Winterbourn MJ & Gregson KLD. 1981. Guide to the aquatic insects of New Zealand. Bulletin 39 40 of the Entomological Society of New Zealand 5. 80 p. 41 42 Winterbourn MJ, Rounick JS & Cowie B. 1981. Are New Zealand stream ecosystems really 43 44 different? New Zealand Journal of Marine and Freshwater Research 15 , 321–328. 45 46 47 Worthy TH, Tennyson AJD, Hand SJ & Scofield RP. 2008. A new species of the diving duck 48 Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in the St Bathans Fauna 49 50 (Early Miocene), New Zealand. Journal of the Royal Society of New Zealand 38 , 97–114. 51 52 53 Worthy TH, Tennyson AJD, Hand SJ, Godthelp H, Scofield RP 2011. Terrestrial turtle fossils 54 55 from New Zealand refloat Moa’s Ark. Copeia 1, 72–76. 56 57 58 59 60 Austral Entomology Page 76 of 81

76 1 2 3 Yavorskaya M, Leschen RAB, Polilov AA, & Beutel RG. 2014. Unique rostrate larvae and 4 5 basidiomycophagy in the beetle family Corylophidae. Arthropod Structure & 6 7 Development 43 , 153–62. 8 9 10 11 12 13 14 15 16 17 18 For Review Only 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 77 of 81 Austral Entomology

77 1 2 3 Figure Captions 4 5 6 7 8 9 Fig. 1. Habitus illustrations of four notable New Zealand insects. (a) Deinacrida rugosa Buller, a 10 giant weta and one of the largest orthopterans in the world. This species was formerly 11 12 widespread on the mainland of North Island, but is now restricted to a few predator free offshore 13 14 island. (b) Chalcodrya variegata Redtenbacher a species from the endemic beetle family 15 16 Chalcodryidae. This species is associated with the sooty mould fauna. (c) Micrarchus nov. sp. 2, 17 an undescribed species from northwest Nelson and Westland that is restricted to high alpine 18 For Review Only 19 areas. Genomic studies are revealing unique adaptations to these harsh conditions. (d) 20 21 Zelandochlus latipalpis Brundin, the 'Ice Worm', a flightless nonbiting midge species known 22 23 from New Zealand's glacier fields. This taxon is thought to have diverged from its sister in the 24 25 Oligocene. Illustrations (a, (b), and (c) by Des Helmore (© Landcare Research). Illustration (d) 26 reproduced from Brundin (1966). 27 28 29 30 31 32 Fig. 2. Habitats containing notable New Zealand insect species. (a) The fastflowing Cleddau 33 River slices a path through Nothofagaceae forest in Fiordland. This river is typical of many New 34 35 Zealand freshwater ecosystems and is inhabited by many endemic aquatic insect groups. (b) 36 37 Chancellor Shelf, Fox Glacier, home to a wide variety of alpine lineages, many showing unique 38 39 adaptations to survive in the alpine zone. (c) Carnley Harbour, as seen from Adams Island in the 40 41 subantarctic Auckland Islands. (d) KanukaBroadleaf forest, Karikari Peninsula, Northland. 42 Many of these volcanic uplands in Northland were once islands and still harbour endemic 43 44 species, witnessing their allopatric past. Photo credits: (a) M. Krosch, (b–d) R. A. B. Leschen. 45 46 47 48 49 Fig. 3. Map of New Zealand with place names mentioned in the text marked. The inset map 50 51 shows the New Zealand region including offshore islands. 52 53 54 55 56 57 58 59 60 Austral Entomology Page 78 of 81

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