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Investigation into New Zealand endemic leaf ( and ) and attempts to locate in Kahurangi National Park for host testing against Eadya daenerys, a potential biocontrol agent

Carl Wardhaugh, Andrew Pugh, Matt Scott, and Toni Withers

Report information sheet

Report title Investigation into New Zealand endemic leaf beetles (Chrysomelinae and Galerucinae) and attempts to locate species in Kahurangi National Park for host testing against Eadya daenerys, a potential biocontrol agent

Authors Carl Wardhaugh, Andrew Pugh, Matt Scott, and Toni Withers, Scion

Client Department of Conservation

SIDNEY output 60686 number

Report Number 23807

Signed off by S. Pawson

Date 29 June 2018

Confidentiality Publically available requirement

Intellectual © New Zealand Forest Research Institute Limited. All rights reserved. Unless property permitted by contract or law, no part of this work may be reproduced, stored or copied in any form or by any means without the express permission of the New Zealand Forest Research Institute Limited (trading as Scion).

Disclaimer The information and opinions provided in the Report have been prepared for the Client and its specified purposes. Accordingly, any person other than the Client uses the information and opinions in this report entirely at its own risk. The Report has been provided in good faith and on the basis that reasonable endeavours have been made to be accurate and not misleading and to exercise reasonable care, skill and judgment in providing such information and opinions.

Neither Scion, nor any of its employees, officers, contractors, agents or other persons acting on its behalf or under its control accepts any responsibility or liability in respect of any information or opinions provided in this Report.

Published by: Scion, 49 Sala Street, Private Bag 3020, Rotorua 3046, New Zealand. www.scionresearch.com

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Executive summary

Introduction Native chrysomeline leaf beetles need to be tested against the braconid parasitoid wasp Eadya daenerys (hereafter referred to as Eadya), a proposed biological control agent for the invasive eucalypt tortoise , charybdis. Eadya attacks the larval stages of diurnal, exposed, leaf-feeding chrysomelines that are active during the early summer (December). At least nine species were identified as high-priority species to test with Eadya (two exotic pests, six exotic weed biocontrol agents, and at least one native species). By 2017 host-range testing with Eadya had been successfully completed for the two pests and six weed biological control agents on the host specificity testing list. Results to date are promising and suggestive that Eadya is host specific to the tribe paropsini, as presented in the Scion Technote: (https://www.scionresearch.com/__data/assets/pdf_file/0014/64004/ParopsisTechNote.pdf ). Arguably representatives of our native chrysomeline fauna are the most important species for host testing, however little is known about the larval stages, or host , of these species.

This project Our aim was to locate and study these beetles to determine their potential vulnerability to Eadya, and to identify sources of beetles for host range testing if needed. The best outcome would be locating a few species for testing, and collecting sufficient larvae that they could be transferred to the containment facility in Rotorua for host testing against Eadya. The first step was to examine all literature and collection records to prioritise species for host testing that may potentially be affected by Eadya. The next step was to undertake field work at the best time of year to find, collect priority species, and then conduct host specificity tests.

Key results From the available information on our native NZ chrysomelid fauna (body size, distribution, larval behaviour, and phylogenetic relationship) we conclude that the vast majority of native species are unlikely to be affected by the introduction of Eadya. Most Chrysomelidae are likely to be too distantly related to attract Eadya or support its development. Of the more closely related species, most are too small to facilitate full physiological development of an Eadya , or live in habitats (high-elevation sites, roots, or leaf litter) where Eadya does not search for potential hosts. We consider that species most closely related to Paropsis (i.e. from the subfamily Chrysomelinae), that feed on (Myrtaceae), that are large enough to potentially support the development of an Eadya larva (>5mm), and that have diurnal, external, leaf-feeding larvae that are present in the late spring/early summer period to be most vulnerable to attack by Eadya. Using these biological and ecological filters, we narrowed our focus to just a few of the 150+ native chrysomelid species. The most likely candidates were Chalcolampra speculifera, Cyrtonogetus crassus, or one of the larger species of Allocharis or Caccomolpus. Historical collections of beetles potentially vulnerable to Eadya attack indicated that several large chrysomeline species were present in Kahurangi National Park. A DoC permit was obtained, and subsequent field searches were made in November and December 2017, and January 2018. We located populations of what we believe to be one of the most appropriate species (Allocharis nr. tarsalis) to test against Eadya, based on its phylogenetic relatedness to Paropsis (same subfamily), its body size (second in size only to the nocturnal, refuge-building Chalcolampra speculifera), and larval biology (daytime

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active, exposed leaf-feeding larvae present in the early summer). Host-testing trials with this species should reveal if Eadya is likely to pose any threat to other native leaf beetle species.

Implications of results for the client

The Department of Conservation and local iwi (Manawhenua ki Mohua) may be interested in this new information on the biology and ecology of a little known species, Allocharis nr. tarsalis. New information includes the identification of its host plant Veronica albicans, and distribution across Mount Arthur and Mount Peel in Kahurangi National Park. This will greatly add to what was previously known about this species, which was largely obtained from label information on pinned adult specimens in the National Collection (NZAC) at Manaaki Whenua Landcare Research, Auckland.

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Investigation into New Zealand endemic leaf beetles (Chrysomelinae and Galerucinae) and attempts to locate species in Kahurangi National Park for host testing against Eadya daenerys, a potential biocontrol agent

Table of contents

Executive summary ...... 3 Introduction ...... 6 Information from the literature ...... 7 Field collection methods ...... 21 Results and discussion ...... 22 Conclusions ...... 27 Acknowledgements ...... 28 References ...... 29 Appendix A ...... 32 Appendix B ...... 33 Appendix C ...... 34 Appendix D ...... 35 Appendix E ...... 36 Appendix F ...... 37

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Introduction

The proposed biological control project New Zealand has ~150 native species of leaf beetles (Coleoptera: Chrysomelidae), while another ~30 species have been accidentally or deliberately introduced (MacFarlane et al. 2010). One of New Zealand’s most damaging exotic pests is the eucalyptus tortoise beetle, Stål. (Chrysomelidae: Chrysomelinae: Paropsina), which has been present for over 100 years and remains the most significant pest of commercial eucalypt plantations (Murphy & Kay 2000; Withers & Peters 2017). The larvae and adults feed on the leaves of a range of Eucalyptus species (Myrtaceae), inhibiting growth and even, on occasion, resulting in mortality (Murray et al. 2008; Murray et al. 2010). Currently, expensive aerial spraying of insecticides is the primary control measure for Paropsis (Withers et al. 2013). Classical biological control agents have been sought to reduce Paropsis impacts on host to mitigate the need for insecticides. To date, several biological controls have been introduced, with limited success (Bain & Kay 1989). A promising biological control agent from Tasmania, Eadya daenerys (Hymenoptera: Braconidae), has been identified, which targets first generation larvae of Paropsis each spring (Rice 2005; Rice & Allen 2009; Peixoto et al. 2018). The spring larva is a particularly vital stage to target as none of the existing agents have a significant impact on this important first seasonal generation (Murphy & Kay 2000; Mansfield et al. 2011). Eadya daenerys 2018 Ridenbaugh (hereafter referred to as Eadya) is a parasitoid wasp that feeds internally on the larvae of tortoise beetles from the closely-related genera Paropsis and (Peixoto et al. 2018). Before any decision can be made regarding the release of Eadya, the potential impacts on non-target species in New Zealand must be assessed (Barratt et al. 2006; Todd et al. 2016). Non-target species were selected based on phylogenetic relatedness and niche overlap with Paropsis (i.e., all closely-related leaf beetle species and those more distantly-related leaf beetles whose larvae have a similar ecology to Paropsis larvae) (Withers et al. 2015; Withers et al. 2018). Eight exotic pest species or weed biological control agents were chosen for host-testing against Eadya in a series of no-choice and two-choice (with Paropsis) behavioural assays and physiological development tests (Withers et al. 2017). Risks to native species are critical to the assessment of potential environmental impacts of a new biological control agent. Therefore, locating and testing native leaf beetles became one of the most important aspects of the project (Withers et al. 2015).

New Zealand endemic Chrysomelidae Our endemic leaf beetle fauna is poorly known, both taxonomically and ecologically, but includes a number of species in the same subfamily as the target pest beetle (see Table 1). As Eadya can attack all larval stages of its paropsine hosts we were most interested in exposed, leaf-feeding non-target beetle larvae that are active during the day in late spring and early summer (Rice 2005). Current knowledge indicates that Eadya is specialized to a small number of host beetle species. It has only been reared from paropsine tortoise beetles in its native range (Tasmania) (Peixoto et al. 2018) and ignores other beetle larvae present on the same leaves at the same time (e.g. Gonipterus platensis) (G. Allen, University of Tasmania, pers. comm). Because of this apparent host specificity in the country of origin, we focus on species from the subfamily Chrysomelinae (to which Paropsis

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belongs), and the sister subfamily Galerucinae (Table 1) (Reid 1995). Other than distant phylogenetic relatedness, all other subfamilies may be discounted based on larval characteristics (see Table 1).

Information from the literature

Sub- Chrysomelinae New Zealand has over 40 native species of Chrysomelinae in 5 genera: Allocharis, Aphilon, Caccomolpus, Chalcolampra and Cyrtonogetus (Leschen & Reid 2004; Reid 2006). All of these species belong to the subtribe Phyllocharina, while the introduced Paropsis belongs to a separate subtribe; Paropsina. Species of Phyllocharina are distributed across Australia, New Caledonia, and South America (Reid & Smith 2004; Reid 2006; Jurado-Rivera et al. 2009; Reid et al. 2009). There is little biological information on these naturally uncommon native species (Leschen et al. 2012), with the exception of some more detailed observations on the following three species: Chalcolampra speculifera, Allocharis robusta, and A. marginata (Hudson 1934; Wardle et al. 1971; Reid 1995; Jolivet & Hawkeswood 1995) The larvae of Chalcolampra speculifera are nocturnal, hiding during the day in refuges bored into the stems of their host plants, colensoi Hook.f. var. colensoi () (Wardle et al. 1971). The entrance hole is plugged with a scleritised anal plate covered in spines; “the apical plate acts as a plug for the diurnal burrows of nocturnal larvae” (cited as J. Dugdale, personal communication in Reid (1995)). Similar larvae in the New Zealand Arthropod Collection (possibly from another undescribed species of Chalcolampra) have been collected from mountain daisies (Celmisia spp.) (Figure 1) in New Zealand. Label information from adult specimens stored at the NZAC shows that most Chalcolampra specimens have been collected only in alpine and sub-alpine habitats, such as the Old Man Range in Otago and on Mount Arthur in Kahurangi National Park, Nelson. Larvae of Allocharis robusta Broun were observed feeding on the leaves and flowers of Veronica (Plantaginaceae) at Aoraki/Mt Cook (Hudson 1934). Hudson (1934) describes these as “fairly active,” dark in colour, and about 9 mm long when fully grown. Hudson’s illustrations (Figure 2) show no sign of an anal plate so these species presumably feed exposed and do not make refuges. It was not mentioned at what time of day they were active. Larvae and adults of Allocharis marginata Broun were reported feeding on Veronica salicifolia along mountain riverbanks (Jolivet & Hawkeswood 1995), though no more specific information was supplied. These scant records indicate that a large species of Allocharis, rather than the nocturnal, refuge-building Chalcolampra, would be an ideal candidate for host testing against Eadya.

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Figure 1: On the left, the larva of an unknown species of Chrysomelinae collected from a mountain daisy, Celmisia sp Asteraceae (NZAC wet collection). This larva may be from an undescribed species of Chalcolampra, or it could be that C. speculifera may attack Celmisia in addition to their current known host plant, . It appears very similar to the drawing on the right from Reid (1995), which despite the label indicating it is a species of Allocharis, we believe is actually a Chalcolampra larva. This assumption is supported by the presence of the sclerotized anal plate used by Chalcolampra speculifera to plug the daytime refugia spoken of by John Dugdale and photographed by Wardle et al. (1971), and the lack of such a structure in the illustrations of A. robusta in Hudson (1934).

Figure 2: Illustration of the black leaf-feeding larva and the adult of Allocharis robusta, collected from Veronica sp. near Mt Cook village in the 1930’s (Hudson 1934).

Australian species of Chalcolampra form two clades based on molecular evidence; the first group feed on host plants in the Order Lamiales, while the second group feed on (C. Reid pers. comm.). It is possible that these two clades are represented by two genera in New Zealand; Chalcolampra that feeds on Asterales, i.e. Olearia and Celmisia and the closely-related Allocharis that feeds on Lamiales, i.e. Veronica. In New Caledonia, another close relative in the Zira is restricted to trees of the family Myodocarpaceae, formerly Araliaceae (Jolivet & Verma 2004; Reid & Smith 2004). Beetle taxonomist Richard Leschen is currently reviewing the of these genera Chalcolampra and Allocharis (R. Leschen pers. comm.) but the results are not available at the time of writing. The larvae of some species in the genus Aphilon (13 spp.) have been collected from mosses and liverworts at night (Kuschel 1990). These minute species are not of interest in this study of parasitoid risks due to their small body size (<3.5mm), nocturnal habits, and ground-based host plants. The closely- related genus Caccomolpus (16 spp.) may also have moss-feeding larvae, based on the fact that many adult beetles have been collected from mosses, under rocks, and in leaf litter (see Tables 2 and 3). Most species in this genus are also likely to be too small (<5mm), for complete development by Eadya even if they were attacked, although the largest species (6mm) could potentially be vulnerable. Adults of Cyrtonogetus crassus have been collected from mountainous areas of Central Otago (Broun 1915) but the larvae are unknown.

Sub-family Galerucinae Galerucinae are a diverse subfamily consisting of more than 11,000 described species, but fewer than 100 are present in New Zealand. Several phylogenetic analyses have shown that Galerucinae are a sister group to Chrysomelinae (Reid 2014). Our native Galerucinae are divided into two tribes; the and the

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Alticini (Nadein & Bezděk 2014). Based on the biology of closely-related species overseas we presume that our native Galerucini have root-feeding larvae (Hua et al. 2014). All Galerucini in New Zealand belong to the subtribe Luperina, which have the common name rootworms because of the feeding biology of the larvae. The larvae of all of our native species are unrecorded, presumably because they feed in this cryptic manner. The Alticini are more varied in their larval biologies (Samuelson 1973). However, all New Zealand species are very small, <4.5mm in length. Eadya is a relatively large wasp that attacks the relatively large larvae of Paropsisterna and Paropsis (~8-12mm long) (Peixoto et al. 2018). Indeed, P. charybdis is the largest species in the family Chrysomelidae in New Zealand. Based on their size, Eadya is highly unlikely to succeed in using native Alticini as hosts, or in fact any host beetle smaller than 8mm long as an adult (Tables 1 and 2). Despite the minimal risk posed by Eadya to any small native Alticini, we included in our host testing list the largest species in this tribe present in New Zealand (the exotic biological control agent hygrophila, 5.05mm long). It was also included because its diurnal larvae feed externally on alligator weed during the spring and summer, so it places it at higher risk of attack than root- feeding native Galerucinae (Stewart et al. 1999).

Table 1: Summary of the diversity of each subfamily of Chrysomelidae in New Zealand, their potential vulnerability to attack by Eadya daenarys if it were released based purely on habitat or phenological overlap with Eadya/Paropsis charybdis populations, and the reasons why they may or may not be vulnerable to a larval parasitoid that searches externally on leaves. Subfamily No. of Vulnerability Reasons spp. in of native spp. NZ to Eadya (native) Chrysomelinae 52(42) Low to Native spp. are more distantly Moderate related than some exotic species to Paropsis. Most species either very small (<4.5mm), or likely to be nocturnal, or present in mosses/liverworts, or a combination. Naturally uncommon. Most spp. restricted to high-elevation sites in the South Island. Some are leaf- feeders. Largest species shows adaptation to living in stem refugia. Galerucinae 83(77) Low/none Relatively closely related, considered a sister group to above. All native species either very small (<4.5mm) or almost certainly develop in the soil, or both. Many spp. restricted to high-elevation sites. Only some exotic species are known leaf miners or external leaf feeders.

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Eumolpinae 19(19) Very low/none Distantly related. Larvae develop in the soil. 10(8) Very low/none Distantly related. Larvae of native species are case-bearers that live in the leaf litter. 4(0) None Distantly related. No native spp. 2(0) None Distantly related. No native spp. Bruchinae 3(0) None Distantly related. Larvae feed internally in seeds. No native spp.

Non-target species selection On the basis of the available biological and ecological information we conclude that the larvae of at least one of our larger species of native chrysomelines should be tested against Eadya in host-testing trials (Withers et al. 2015). Criteria for selecting appropriate species are: large enough to be of interest to Eadya (>5mm), external leaf-feeding larvae, diurnal activity, and larvae present during the spring/summer flight period of Eadya. Based on what little information we have on our native Chrysomelinae, the most likely beetle to meet these criteria would therefore be the large Allocharis species, such as A. tarsalis or A. robusta (see Table 2).

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Table 2: List of all Chrysomelinae and Galerucinae species present in New Zealand. Letters after the species names indicate the origin of the species (A = Adventive, E = Endemic, BC = Biological Control). The average length of each species is given. This information has been obtained either from type specimens listed in the literature, or by directly measuring specimens in collections. The elevation at which a species lives, its host plants, and habitat preferences are indicated where these are known. Size Elevation Subfamily/(Sub)tribe/Species (mm) (m) Habitat/host plants/notes References Chrysomelinae Chrysolinina abchasica (Weise 1892) A BC 5.5 Tutsan (Förster, 1771) A BC 6 St. John's wort (Groenteman et al. 2011) (Suffrian, 1851) A BC 7 St. John's wort Dicranosternina Dicranosterna semipunctata (Chapuis, 1877) A 9 Acacia trees (Murray & Withers 2011) Gonioctenina (Spartophila) olivacea (Förster, 1771) 5 Scotch broom (Syrett 1989) BC Paropsina Paropsis charybdis Stål, 1860 A 11 Lowland Eucalypt plantations (Kuschel 1990) Paropsisterna beata (Newman, 1842) A 11 Lowland Eucalypt plantations (Yamoah et al. 2016) Paropsisterna variicollis (Chapuis, 1877) A 8 Lowland Eucalypt plantations (Lin et al. 2017) Pyrgoides sp. 6 A Acacia trees (Kuschel 1990) catenata (Chapuis, 1877) A 8.5 Lowland Eucalypt plantations (Barrett 1998) (Blackburn, 1896) A 8 Lowland Eucalypt plantations (Simmul & De Little 1999) Phyllocharina Allocharis fuscipes Broun, 1917 E 4.5 600 Moa Basin, Wilberforce river (Broun 1917) Allocharis limbata Broun, 1893 E 4.5 600 Flagstaff Hill, Dunedin, Mt. Maungatua, (Broun 1893) Dansey's Pass, mat plants Allocharis marginata Sharp, 1882 E 5.5 Craigieburn, Mt Hutt, mat plants Ref Order

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Allocharis media Broun, 1917 E 4.5 Ben Lomond, Mt Earnslaw (Broun 1917) Allocharis morosa Broun, 1893 E 5 (Broun 1893) Allocharis nigricollis Broun, 1917 E 5 1300 Ben Lomond, Mt Dick (Broun 1917) Allocharis picticornis Broun, 1917 E 4.5 Ben Lomond (Broun 1917) Allocharis praestans Broun, 1917 E 5.5 Moa basin (Broun 1917) Allocharis robusta Broun, 1917 E 6 Lake Wakatipu, Aoraki Mt Cook (Broun 1917) Allocharis subsulcata Broun, 1917 E 5 1300 Old Man Range, Veronica odora, Celmisia (Broun 1917) prorepens Allocharis tarsalis Broun, 1917 E 7 Gordon's Knob, Belgrove, Richmond FP (Broun 1917) Aphilon convexum Broun, 1893 E 2 Howick, ground vegetation (Broun 1893) Aphilon enigma Sharp, 1876 E 2 Auckland, moss and liverworts Aphilon impressum Broun, 1914 E 3.5 McClennan's bush, Methven, amongst leaf (Broun 1914) mould Aphilon laticolle Broun, 1893 E 3.5 Thames (Broun 1893) Aphilon latulum Broun, 1893 E 2 Stratford, on a log (Broun 1893) Aphilon minutum Broun, 1880 E 2 Whangarei, Auckland, moss and liverworts (Broun 1880) Aphilon monstrosum Broun,1886 E 3 Waitakerei ranges, Auckland, moss and (Broun 1886) liverworts Aphilon praestans Broun, 1893 E 2.5 (Broun 1893) Aphilon pretiosum Broun, 1880 E 3.5 Tairua, Auckland (Broun 1880) Aphilon punctatum Broun, 1880 E 3.5 Whangarei (Broun 1880) Aphilon scutellare Broun, 1893 E (Broun 1893) Aphilon sobrinum Broun,1886 E 3 Waitakerei ranges (Broun 1886) Aphilon sternale Broun, 1921 E 3.5 Auckland (Broun 1921) Caccomolpus amplus Broun, 1921 E 6 200-1000 Glenhope, Mt Dewar, moss at night (Broun 1921) Caccomolpus cinctiger Broun, 1921 E 4.5 Glenhope (Broun 1921) Caccomolpus flectipes Broun, 1914 E 5 1100 Mt Hutt (Broun 1914) Caccomolpus fuscicornis Broun, 1917 E 4.5 Mt Dick, Stewart Island to Nelson (Broun 1917) Caccomolpus globosus Sharp, 1886 E 3 Greymouth, Fiordland (Broun 1886) Caccomolpus hallianus Broun, 1917 E 5 Mt Dick, Lake Wakatipu, decaying leaves (Broun 1917) Caccomolpus maculatus Broun, 1893 E 4 Mt Arthur (Broun 1893)

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Caccomolpus montanus Broun, 1921 E 5 Mt St. Arnaud (Broun 1921) Caccomolpus nigristernis Broun, 1917 E 3 Hollyford, amongst decaying vegetation (Broun 1917) Caccomolpus ornatus Broun, 1910 E 3.5 Waimarino (Broun 1910) Caccomolpus plagiatus Sharp, 1886 E 4 Greymouth Caccomolpus pullatus Broun, 1893 E 4 Forty Mile Bush (Broun 1893) Caccomolpus subcupreus Broun, 1921 E 5 Glenhope (Broun 1921) Caccomolpus substriatus Broun, 1917 E 4.5 Ben Lomond, decaying vegetation (Broun 1917) Caccomolpus tibialis Broun, 1917 E 4 Mt Dick (Broun 1917) Caccomolpus viridescens Broun, 1917 E 5.5 Dyer's Pass, Christchurch (Broun 1917) Chalcolampra (Eualema) speculifera Sharp, 1882 E 8 Sealevel to 1300m, most records >500m. (Broun 1910) Olearia colensoi Cyrtonogetus crassus Broun, 1915 E 6.5 Remarkables (Broun 1915) Galerucinae Luperina Adoxia aenea Broun, 1880 E 4 Lindis Pass (Broun 1880) Adoxia aenescens (Sharp, 1886) E 4 Bealey (Broun 1886) Adoxia angularia (Broun, 1909) E 5.5 Broken River (Broun 1909) Adoxia anthracina (Broun, 1914) E 3 1100 Hump Ridge (Broun 1914) Adoxia asperella (Broun, 1909) E 7 South Island (Broun 1909) Adoxia atripennis (Broun, 1913) E 6.5 Lake Wakatipu (Broun 1914) Adoxia attenuata Broun, 1880 E 5 Whangarei (Broun 1880) Adoxia aurella (Broun, 1914) E 4.5 Mt Hutt (Broun 1914) Adoxia axyrocharis (Broun, 1909) E 3.5 Arthur's Pass (Broun 1909) Adoxia brevicollis (Broun, 1893) E 5 Mt Egmont (Broun 1893) Adoxia bullata (Broun, 1914) E 4.5 1350 Mt Hutt (Broun 1914) Adoxia calcarata (Broun, 1893) E 5 Mt Arthur (Broun 1893) Adoxia cheesemani (Broun, 1910) E 3.5 Mt Cook (Broun 1910) Adoxia cyanescens (Broun, 1917) E 5 Moa Basin (Broun 1917) Adoxia dilatata (Broun, 1914) E 4.5 Mt Hutt (Broun 1914) Adoxia dilucida (Broun, 1917) E 5.5 Moa Basin (Broun 1917)

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Adoxia dilutipes (Broun, 1915) E 3.5 Ben Lomond (Broun 1915) Adoxia discrepans (Broun, 1914) E 6.5 Mt Dennan, Tararua range (Broun 1914) Adoxia diversa (Broun, 1910) E 3 Mt Cook (Broun 1910) Adoxia foveigera (Broun, 1913) E 6.5 Capleston, Westland (Broun 1914) Adoxia fuscata (Broun, 1893) E 4 1000 Mt Egmont (Broun 1893) Adoxia fuscifrons (Broun, 1910) E Adoxia gracilipes (Broun, 1917) E 4.5 Moa Basin (Broun 1917) Adoxia halli (Broun, 1917) E 5.5 Scarcliff, Moa Hill (Broun 1917) Adoxia insolita (Broun, 1914) E 4 1100 Hump Ridge (Broun 1914) Adoxia iridescens (Broun, 1914) E 4 Mt Hutt (Broun 1914) Adoxia lewisi (Broun, 1909) E 5 Broken River (Broun 1909) Adoxia mediocris (Broun, 1917) E 4.5 Moa Basin (Broun 1917) Adoxia minor (Broun, 1917) E 3.5 Moa Basin (Broun 1917) Adoxia mollis (Broun, 1893) E Adoxia monticola (Broun, 1893) E 4 Mt Maungatua, Taieri (Broun 1893) Adoxia nigricans Broun, 1880 E 4.5 Tairua (Broun 1880) Adoxia nigricornis (Sharp, 1886) E 5 Greymouth (Broun 1886) Adoxia nitidicollis Broun, 1880 E 3.5 Lindis Pass, Oamaru (Broun 1880) Adoxia nodicollis (Broun, 1915) E 6.5 Ben Lomond (Broun 1915) Adoxia obscura (Broun, 1910) E 3.5 Mt Cook (Broun 1910) Adoxia oconnori (Broun, 1913) E 5 Ohau, near Wellington (Broun 1913) Adoxia oleareae (Broun, 1893) E 5 1100 Mt. Egmont, on Olearia nitida (Broun 1893) Adoxia palialis (Broun, 1909) E 4 Broken River (Broun 1909) Adoxia perplexa (Broun, 1917) E 6 Scarcliff, Moa Basin, Broken River (Broun 1917) Adoxia princeps (Broun, 1893) E 5 1800 Mt Tyndall (Broun 1893) Adoxia proletaria (Weise, 1924) E Adoxia pubicollis (Broun, 1915) E 3.5 Takitimu mountains (Broun 1915) Adoxia puncticollis (Sharp, 1886) E 4.5 Otira, Wakefield, Kowhai trees Adoxia pygidialis (Broun, 1917) E 3.5 Lake Wakatipu (Broun 1917) Adoxia quadricollis (Broun, 1917) E 3 Moa Basin (Broun 1917)

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Adoxia rectipes (Broun, 1893) E 5 Otira Gorge (Broun 1893) Adoxia rugicollis (Broun, 1893) E 4.5 1100 Mt Egmont (Broun 1893) Adoxia scutellaris (Broun, 1909) E 5.5 Broken River (Broun 1909) Adoxia simmondsi (Broun, 1913) E 5 1200 Mt Quoin, Tararua Range, on Olearia colensoi (Broun 1914) Adoxia sordidula (Weise, 1924) E 4 1100 Mt Egmont Adoxia sulcifera (Broun, 1893) E 5.5 Otira Gorge (Broun 1893) Adoxia thoracica (Broun, 1880) E 5 Tairua (Broun 1880) Adoxia truncata (Broun, 1893) E 4 Otira Gorge (Broun 1893) Adoxia vestitus (Weise, 1924) E Adoxia vilis (Weise, 1924) E Adoxia viridis Broun, 1880 E 5 Lindis Pass (Broun 1880) Adoxia vulgaris Broun, 1880 E 5 On Brachyglottus repanda and (Broun 1880) Adoxia xenoscelis (Broun, 1917) E 3.5 Moa Basin (Broun 1917) Allastena eminens Broun, 1917 E 3 Moa Basin (Broun 1917) Allastena nitida Broun, 1893 E 3 Mt Maungatua (Broun 1893) Allastena piliventris Broun, 1915 E 3 Ben Lomond (Broun 1915) Allastena quadrata Broun, 1893 E 3 Mt Maungatua, Otira Gorge (Broun 1893) Bryobates aeratus Broun, 1914 E 4.5 1100 Hump Ridge, Okaka (Broun 1914) Bryobates coniformis Broun, 1886 E 3.5 Mt Maungatua, on moss (Broun 1886) Bryobates nigricans Broun, 1914 E 5 Amongst moss, Clutha (Broun 1914) Bryobates rugidorsis Broun, 1917 E 4 Remarkables (Broun 1917) suturalis Thomson, 1866 BC 5.5 Heather (Syrett et al. 2000) Alticini Selman & Vogt, 1971 BC 5 Alligator weed (Stewart et al. 1999) Alema paradoxa Sharp, 1876 E 3 Auckland, Tairua (Samuelson 1973) Alema spatiosa Broun, 1880 E 4.5 Whangarei (Broun 1880; Samuelson 1973) carduorum (Guérin-Méneville, 1858) BC 4 Canadian thistle (Samuelson 1973) (Chaetocnema) paspalae (Broun, 3 Lake Ohia, Northland Samuelson 1973, 1923) E (Broun 1923)

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Chaetocnema (Tlanoma) aotearoa Samuelson, 1.5 South Island, moss in tussock (Broun 1923; 1973 E Samuelson 1973) Chaetocnema (T.) graminicola (Broun, 1893) E 1.5 Iron hills, grass, mat plants (Broun 1893; Samuelson 1973) Chaetocnema (T.) littoralis (Broun, 1893) E 1.7 South Island Samuelson 1973, (Broun 1893) Chaetocnema (T.) moriori Samuelson, 1973 E Ch 2.4 Chatham Islands, Myositidium hostensia (Samuelson 1973) Chaetocnema (T.) nitida (Broun, 1880) E 2.2 Otago, moss, mat plants, lichens (Samuelson 1973), (Broun 1880) fuliginosus (Broun, 1880) E 2.2 Grass (Samuelson 1973), (Broun 1880) Longitarsus jacobaeae (Waterhouse, 1858) BC 3 Tansy ragwort (Samuelson 1973) undulata (Kutschera, 1860) A 2 Brassicas (Samuelson 1973) Pleuraltica cyanea (Broun, 1880) E 4 Tairua, karamu (Samuelson 1973), (Broun 1880) brettinghami Baly, 1862 A 2.8 Various plants (Samuelson 1973) Trachytetra robusta Broun, 1923 E 2.5 Pokako, in leaf mould (Samuelson 1973), (Broun 1923) Trachytetra rugulosa (Broun, 1880) E 2.2 Tairua (Samuelson 1973), (Broun 1880)

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Table 3: Summary of information on native Chrysomelinae species, including sampling localities, host plants, elevation, seasonality, and body size as taken from examining specimens at NZAC, Landcare Research, Tamaki Campus, Auckland.

Species Body size Altitude/seasonality Locality/habitat Host plant/notes Chalcolampra 3.2-5.2 x 6.0-9.8 0 – 1300m South Island Larvae bore into speculifera Nov-Feb (Routeburn track, elongated shoots Southland; Takaka of Olearia Hill, Waikoropupu colensoi. They Springs, have a modified Whanamoa, Mt anal plate to Arthur, Mt Owen, cover hole. Dun Mt., “Canan”, Pseudopanax Nelson; Arthurs arboreus listed Pass, Canterbury but probably incorrect host. Allocharis 3.1–3.4 x 5.5–5.8 1370m Old Man Range, Celmisia subsulcata Jan Otago prorepens, Veronica, Parahebe, Chalmisia spp. among moss.

Allocharis 2-3.6 x 5-6.4mm 840m Flora Hut, Mt , (undescribed sp.) Nov-Dec Arthur, Mt Dewar, Dracophyllum Westcoast/Nelson. longifolium, Also Old Man Olearia colensoi Range, Mt Peel, Mt Domett, Paparoa Range, Lochnagar Ridge, South Island

Allocharis robusta 3.3 x 6.5mm 1220m Titan Huts, Garvie Veronica sp. Jan Mts, Southland, Mt. Cook.

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Allocharis 600m Wilberforce River, fuscipes Canterbury Allocharis limbata 600m Mt Algidus, Mat plants Canterbury; Dansey’s Pass, Mt Teviot, Waipori, Otago Allocharis 1400m (base of Mt Mt Hutt, Mat plants marginata Hutt) Feb Canterbury Allocharis media 2.0 x 4.3 Jan Mt Earnslaw, Otago

Allocharis 500m Mt Domett, nigricollis Mar Nelson; Ben Lomond, Mataura Valley, Southland Allocharis Oct Moa Basin, praestans Scarcliff Allocharis tarsalis 3 x 7mm 1500m Mt. Arthur, Beating hebes at November 1969 Trilobite Hut, Cobb night; under Valley, Nelson stones and in leaf litter.

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Allocharis near 3 x 6.5mm Nov Buckland Peaks, In Celmisia sp. crassus Paparoa Ranges, swards mosses Nelson and lichens

Caccomolpus 2.8-3.1 x 3.6-4.3 200-1000m? Nov- Mt Dewar, Mosses and ferns amplus Jan Paparoa Range, (probably a small Flora Creek, creeping Nelson; Waiho, Blechnum) Ross, Franz Josef, Westland

Caccomolpus 3.7-4.0 x 5.6-6.2 800-1500m? Flora Hut, Flora Moss front brouni Nov-Feb Creek, Mt Arthur, rockface, Mt Domett, Harwood track Canaan Downs, Takaka Hill, Nelson

Caccomolpus sp. 4mm long Apr 2014 Maungatautari, Collected in a Waikato pitfall trap

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Caccomolpus 3 x 6mm R. Leschen cinctiger labelled these “not cinctiger”

Caccomolpus 2 x 3mm Fiordland Takahe globosus Valley

Caccomolpus 3.3 x 4.4 halianus Caccomolpus 2.6 x 4.0 Stewart Island; fuscicornis South Island, Southland, Central Otago, Nelson Aphilon 1.5 x 1.9 <100 Lynfield, Auckland Mosses and monstosum liverworts. Largest species in genus Aphilon minutum 1.3 x 1.8 <100 Lynfield, Auckland Mosses and liverworts

Aphilon enigma Smaller than A. minutum <100 Lynfield, Auckland Mosses and liverworts

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Field collection methods

Northwest Nelson is a biological hotspot for NZ Chrysomelinae. Several species of interest have been collected (Table 3), including Chalcolampra speculifera, the largest species of Allocharis (A. tarsalis), another large, undescribed Allocharis sp., and the largest species of Caccomolpus (C. amplus). We noted the plant species these beetles had previously been collected from (e.g., Olearia colensoi, Olearia lacunosa, various Celmisia spp., Parahebe spp., and Veronica spp.; see Figure 3 and Table 2), and established a plan to search some of the most accessible areas in that region. We obtained permission from the Department of Conservation (Authorization number 54216-RES; see Appendix A) to sample the Cobb Valley and Mt Arthur areas of Kahurangi National Park and the Takaka Hill Scenic Reserve. Within the permitted area, we established a general search envelope comprising a range of geographic environments and vegetation types from which to choose smaller, more concentrated search areas (Appendices B–E).

a) b) c)

Figure 3: Known locations for a) Veronica albicans, b) Olearia colensoi, and c) Olearia lacunosa in NW Nelson. Each of these species is known, or suspected, to be a host plant for at least one large species of native chrysomeline leaf beetle. Source; the New Zealand Plant Conservation Network: www.nzpcn.org.nz/plant_distrbution_results (accessed June 6, 2018)

We made three field trips to the region from November 2017 to January 2018. This period matches the previous collection dates for our species of interest and overlaps the known flight period of Eadya. We also assumed that native Chrysomelinae in low alpine and subalpine environments may feed only on the fresh early season growth, and undergo one larval generation per annum. Hence, early summer sampling seemed most likely to locate target larvae. On each field trip, we covered a portion of the search envelope, targeting specific habitats where known host plants (e.g., Veronica albicans, Olearia colensoi, Olearia lacunosa, Celmisia spp.) or close relative species were present or abundant. Where we found an abundance of one or more of the host plant species, signs of insect damage, or live Chrysomelinae, we made a concentrated search of the plants in the area. We carefully hand searched all known host species, as well as other species (e.g., Dracophyllum spp., Brachglottis spp., Coprosma spp., Olearia spp., Hoheria spp.). Representative plants at each site were sampled using foliage beating techniques to increase the sampling scope. Some sites were re-visited at night. Potential beetle damage was photographed, and promising specimens were collected for further identification in the laboratory. Unusual swellings on plant parts were also dissected in case these were induced by the feeding habits of native chrysomelines, especially on Olearia colensoi and

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Celmisia sp. since the larvae of Chalcolampra are known to induce such swellings when they bore their daytime refuges into the stems. Appendices B–E show areas where concentrated searches were conducted.

Field Trips 1. The first field trip occurred from November 12-17, 2017 during fine weather and covered the Cobb Valley (Cobb River DoC campsite, Cobb Dam Look- out, Trilobite Hut, Myttons Hut, Cobb Track to Cobb Hut, Round Lake, Fenella Hut; see Appendices B and C), and along the roadside on Takaka Hill (see Appendix D). Search effort was approximately 4 person-days. 2. The second field trip occurred from December 11-15, 2017 during fine weather and covered extensive sectors around the Mt Arthur area (around Flora Hut, along the track up Lodestone, up Mt Arthur, Horseshoe Basin, and the Ellis Basin route; see Appendix E). Search effort was approximately 10 person-days. 3. The third field trip occurred from January 8-11, 2018 in cloudy and wet weather and was carried out in the Cobb Valley/Mt Peel area (around Trilobite Hut and the Cobb River, along the track up Mt Peel, below Lake Peel, extensively around Balloon Hut; see Appendix B), two sites on Takaka Hill (see Appendix D). Search effort was approximately 6 person- days.

Results and discussion

A total of 20 person-days search effort, spread across ~20 square kilometres was undertaken in Kahurangi National Park and Takaka Scenic Reserve in 2017-18. No chrysomeline adults or larvae were found in the first search in November. On our second field trip in December 2017, we found the larvae of a chrysomeline species on Veronica albicans (Pétrie) Cockayne (host identified by M. Scott, Scion). The same chyrsomeline was occasionally found on Veronica topiaria (L.B.Moore) Garn.-Jones (host identified by E. M. Miller, National Forestry Herbarium) where both plants were growing adjacent to each other (Figure 4). Live larvae were collected from seven sites between 1200 and 1400m above sea level on Mt Arthur and Lodestone (Figure 5) (indicated with stars in Appendix E, GPS locations Appendix F). What were likely to be members of the same species were also found on our third field trip in January 2018 in the Mt Peel area, although only 20 larvae were found across two sites on this trip (see Appendix B). Extensive feeding damage on host plants suggested that the larval period was nearing its end by early January. In addition to the relatively large number of larvae we found, we also collected two adult Allocharis, which were later identified as Allocharis nr. tarsalis. These were collected by beating some V. albicans bushes on Mt Arthur (site 3) at night. A single adult of a large species of Caccomolpus was also collected (also from V. albicans at site 3). These three individuals (two A. nr. tarsalis and one Caccomolpus) represent the only adult chrysomelines we found on the three fieldtrips (Figure 6).

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Figure 4. Chrysomelinae larvae were found on Veronica albicans (forefront) and also Veronica topiaria (background) when growing sympatrically like this.

Figure 5. Typical habitat where larvae of Allocharis nr. tarsalis were collected in the vicinity of Mt. Arthur and Mt. Peel, Kahurangi National Park

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Allocharis nr. tarsalis Adult beetles (Figure 6) that developed from our live collections were subsequently identified by Dr. Rich Leschen from Landcare as Allocharis nr. tarsalis Broun, which matched the identification of the two adult Allocharis collected from the same plants. Allocharis tarsalis was our number one species of interest as it is the largest definitively day active member of its genus. As suspected (based on the larval biology of the closely-related species A. robusta) A. nr. tarsalis was found feeding externally on the leaves of a Veronica. Approximately 150 larvae in total were transported on V. albicans cuttings to the containment facility at Scion, Rotorua, for host testing purposes and further study. They were reared on fresh cuttings of V. albicans obtained from Titoki Nursery, 26 Palmer Road, Brightwater, Nelson that had been grown from seeds sourced from Kahurangi National Park and had not previously been sprayed with any insecticides. The first plants were later supplemented by more insecticide-free V. albicans of unknown seed source purchased from Kereru Gardens, Tauranga.

Figure 6: Left, Adult of Allocharis nr. tarsalis reared from field-collected larvae. Actual length 7mm. On the right, the single Caccomolpus collected from site 3 (length = 4mm).

The larvae of A. nr. tarsalis were black in colour and quite large, with mature larvae reaching 10mm in length (Figure 7). They fed on the leaf by scraping away the surface layers, which resulted in dark scars that superficially resemble the dark larvae as they sit on the leaf (Figure 8). We speculate that this feeding behavior could function to camouflage the larvae on the leaves, perhaps offering some measure of protection from visual-hunting predators, such as birds. Indeed, the first larvae we discovered were on a plant where some riflemen (Acanthisitta chloris) had just been foraging. In quarantine, 90% (72/80) of the larvae we monitored reached pupation (Figure 9), which included those exposed to Eadya (results presented elsewhere, T. Withers unpublished data). Only two individuals died of unknown causes, and no natural parasitism was observed, either among the 80 individuals used in host testing trials, or the remaining 70 larvae reared in the laboratory to quantify basic biological parameters. The larval period is estimated to last about one month, based on the fact that all of the larvae we reared in captivity (at a constant 18˚C) reached pupation within three weeks of being collected. We did not collect any eggs or first instar larvae, so the smallest larvae we collected were likely second instars at the least.

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Figure 7: A fully grown Allocharis nr. tarsalis larva, with length and width shown. This particular specimen weighed 26mg.

Figure 8: Feeding damage caused by A. nr. tarsalis larvae to Veronica albicans leaves. Note the close resemblance of the larvae (there are at least five in the left-hand photo) to the feeding damage they cause to the leaf surface.

Figure 9: Pupa of Allocharis nr. tarsalis showing length and width.

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When larvae reached maturity, they dropped to the floor of their containers to pupate amongst the sphagnum moss we placed in each container to mimic a ground layer and maintain higher humidity. This suggests that wild Allocharis larvae drop to the ground as pre-pupae and then pupate amongst leaf litter under their host plants. The pupae lost their black colouration and were pale yellow. Pupation lasted one to two weeks, with newly emerged adult beetles taking another week or two for their cuticle to harden up and darken. Weights are shown for each life history stage (Table 4). The larvae in this table are all in their 4th (final) instar. As a comparison in relation to the biological control project, the weight of a mature Allocharis larva is approximately one eighth that of a fourth instar Paropsis charybdis larva (Table 4). Adult Allocharis nr. tarsalis beetles fed on Veronica foliage, and also flowers. They were strictly nocturnal in their feeding activities, spending the day hiding amongst the sphagnum moss placed on the floor of their containers. This behaviour may explain the previous collection of adult beetles from under stones, or by sifting leaf litter (Table 3). Adult beetles were also quite cold-tolerant, and survived overnight freezing at -20°C. They also remained active when placed in a fridge at 4°C, even reproducing at that temperature. This is obviously an adaptation to the cold temperatures experienced in their subalpine habitat on a nightly basis. In April and May 2018, some of our captive Allocharis laid eggs and produced a small number of larvae. We did not discover these until the larvae had reached second instar, so are still unable to describe the eggs or newly emerged larvae. We do not know if they would naturally produce a second generation in autumn, or if the constant temperature and humidity conditions in the laboratory upset the normal reproductive cycle of these beetles, inducing them to breed when they ordinarily would not.

Table 4: A comparison of the mean fresh weights in mg (range) of the target pest Paropsis charybdis and the native beetle Allocharis nr. tarsalis larvae, prepupae, pupae and adults. Larvae (mature) Prepupa Pupa Adult Male 150 169.4 163.7 153.4 (110-170) Paropsis charybdis (128-225) (125-214) (113-200) Female 230 (140-270) 22.2 22.3 22.1 22.7 Allocharis nr. tarsalis (15-27) (12-29) (16-28) (18-29)

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Conclusions

A review of the taxonomy and biology of our native chrysomelids and that of the proposed biological control agent, Eadya daenerys, suggests that this parasitoid poses little, if any, risk to our native chrysomelid fauna. This risk will be assessed by host testing on of our largest day active, native chrysomelid A. nr. tarsalis (T. Withers, unpublished data). Most native chrysomelid species are too distantly related (Criocerinae, Cassidinae, Bruchinae) or too small (Alticinae, many species from other subfamilies, Tables 1 and 2) to support the physiological development of Eadya. A large proportion of native species inhabit the leaf litter or soil as larvae (Cryptocephalinae, , most Galerucinae, and many native Chrysomelinae) and are unlikely to be encountered by Eadya. Of the few species with daytime active, exposed leaf-feeding larvae (Allocharis spp. may prove to be the only native fauna with this feeding habit) we are confident that we have located the largest, and therefore probably the most suitable, species for host testing. Host testing trials with A. nr. tarsalis should give a strong indication of the maximum potential vulnerability of our native chrysomelids to Eadya. Another characteristic of our native chrysomelid fauna that would afford them an extra level of protection from Eadya is their distribution. A high proportion of species live in subalpine or alpine habitats, particularly species from the potentially vulnerable Chrysomelinae and Galerucinae. Almost all of our native Chrysomelinae larger than 5mm in size have been collected predominantly, or exclusively, from higher elevations (e.g., >800m.a.s.l.) (Tables 1 and 2). We suspect the single record of Chalcolampra speculifera from Waikoropupu Springs mentioned in Table 3 is erroneous as known host plants of Chalcolampra spp. (Olearia colensoi or Celmisia spp.) are not known from this site and all other records are from high-elevation sites. In its native Tasmania, E. daenerys has not been recorded from any sites higher than Moina at 600m a.s.l. It is likely that their host paropsine species, and/or the Eucalyptus species on which paropsines feed do not occur at these higher elevations. The lack of Eucalyptus plantations in subalpine or alpine areas in New Zealand strongly suggests that Eadya would rarely, if ever, move into these areas in search of hosts, making it unlikely to encounter a large native chrysomelid larva in daytime. The most risky scenario would be for one or a small number of Eadya to be blown into the subalpine zone from eucalypt plantations in the lowlands. In this case, a female Eadya may stumble upon the larvae of a native chrysomelid, however the probability of this occurring is further reduced by the patchy distribution of these beetles. Host- testing results will reveal if Eadya are attracted to, or attack, an Allocharis larvae, and whether it would be possible for Eadya to form a self-sustaining population on these native species. Lastly, parasitoid wasps often use chemical cues emitted by host trees to locate areas where they are likely to encounter their target prey. Furthermore, observations in the laboratory show that Eadya is most active (walking around, waving antennae) only when in close proximity to Paropsis larvae. In the presence of other species, Eadya would typically remain motionless, or groom itself. These observations indicate that Eadya utililses chemical cues to locate their host targets. It is unknown what host plant volatiles Eadya specifically uses to locate suitable paropsine habitat, but there is no evidence that Veronica spp. (Plantaginaceae) have a similar chemical profile to Eucalyptus (Myrtaceae).

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Host testing results (T. Withers, unpub. data) are available on this website link: https://www.scionresearch.com/__data/assets/pdf_file/0014/64004/ParopsisTechN ote.pdf

Acknowledgements

Funding for this project was provided by the MPI Sustainable Farming Fund (CO407964), NZ Farm Forestry Association, Southwood Exports Ltd, Oji Fibre Solutions NZ Ltd and Scion MBIE Strategic Science Investment funding. Voucher specimens of Allocharis nr tarsalis will be deposited in the New Zealand Arthropod (NZAC) Collection and additional specimens at Scion FRNZ. Collections at Kahurangi National Park were made under DOC research authority 54216RES. We are grateful for the assistance and/or advice of John Dugdale, Richard Leschen (Manaaki Whenua Landcare Research), Greg Napp (DoC), Bev Purdie (Manawhenua ki Mohua), Belinda Gresham (Scion) and Chris Reid (University of Sydney).

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Appendix A

DOC research authority 54216RES.

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Appendix B

Sites visited during field trips 1 and 3, Cobb Valley and Mt Peel surrounds, Kahurangi National Park. Highlighted areas indicate intensive search of host plant patches. Stars indicate positive finds of Allocharis nr tarsalis (Coleoptera: Chrysomelidae: Chrysomelinae)

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Appendix C

Sites visited during field trip 1, Cobb Valley, Kahurangi National Park. Highlighted areas indicate intensive search of host plant patches.

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Appendix D

Sites visited during field trips 1 and 3, Takaka Hill Scenic Reserve and roadsides. Highlighted areas indicate intensive search of host plant patches.

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Appendix E

Sites visited during field trip 2, Mt Arthur and Lodestone vicinity, Kahurangi National Park. Highlighted areas indicate intensive search of host plant patches. Stars indicate positive finds of Allocharis nr tarsalis (Coleoptera: Chrysomelidae: Chrysomelinae).

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Appendix F

Co-ordinates and elevations of sites (indicated by stars in Appendices B and F) where Allocharis nr. tarsalis was collected.

Location Site Elevation number Latitude Longitude (m) Mt Arthur SITE1 -41.1715 172.7429 1254 Mt Arthur SITE3 -41.1977 172.7133 1346 Mt Arthur SITE3 END -41.1982 172.7100 1335 Mt Arthur SITE7 -41.2035 172.6924 1349 Mt Arthur SITE8 -41.2004 172.6921 1355 Mt Arthur SITE9 -41.2014 172.6914 1367 Mt Arthur SITE11 -41.2289 172.6875 1394 Mt Arthur SITE12 -41.1975 172.7155 1296 Balloon Hut B1 -41.1679 172.2651 1225 Balloon Hut B2 -41.1679 172.6236 1245

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