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Home , Aphelinidae, Hyperomyzus, Therioaphis, Therioaphis trifolii

EXOTIC INTRODUCTIONS OF PRIMARY OF IN NEW ZEALAND: THE GOOD AND THE BAD

D.A.J. Teulon, G.M. Drayton, & I.A.W. Scott

New Zealand Institute for Crop & Food Research, Private Bag 4704, Christchurch, New Zealand, [email protected]

ABSTRACT.

This paper summarises the primary species (a mixture of intentionally introduced species, self-introduced species and probably indigenous species) recorded from introduced and indigenous aphids in New Zealand. It reviews the effectiveness of the intentionally and self-introduced primary parasitoids on aphid pests in New Zealand in terms of their ability to control pest aphid species and contrasts this with their potential impact on indigenous aphids.

The majority of aphids found in New Zealand have been inadvertently introduced (about 100 species), including many that are important plant pests in horticulture, agriculture, and forestry. In recent years a growing number of indigenous New Zealand aphid species (currently over 15) have been recognised that are of considerable scientific interest. Most of these indigenous species were discovered after the biocontrol introductions for pest species.

Of the primary parasitoid species purposefully introduced, several are considered to be effective biological control agents, e.g. Aphidius rhopalosiphi and Aphidius ervi. Additionally, other self-introduced species are also considered to be effective biological control agents, e.g. Aphidius colemani. There is increasing evidence that several intentionally and self-introduced primary parasitoid species are attacking indigenous aphid species in New Zealand and we are endeavouring to quantify their impact on these species. The lack of documented morphological characteristics for some species and the presence of undescribed indigenous genera/species in New Zealand make primary aphid parasitoid identification very difficult. The use of molecular techniques has in some part improved the reliability of identification.

INTRODUCTION.

Invasive aphids.

The majority of aphids found in New Zealand have been inadvertently introduced (about 100 species) (Teulon & Stufkens 2002). They have a significant economic impact on arable, vegetable, fruit and ornamental crops, and forest trees, and have unquantified impacts on natural systems. Aspects of their biology, such as their small size, parthenogenetic reproduction, high reproductive rate, short generation time, rapid dispersal and eruptive population dynamics pose difficult challenges for biosecurity and pest management in New Zealand (Teulon & Stufkens 2002).

Procceedings of the Third International Symposium on Biological Control of , Christchurch, New Zealand. Peter G. Mason, David R. Gillespie & Charles Vincent (2008) Proceedings of ISBCA 3 – P. G. Mason, D. R. Gillespie and C. Vincent Eds. (2008)

Indigenous aphids.

Over 15 species of native aphids have also been recorded in New Zealand, with the majority of these being recognised in the last 10 years (Teulon et al. 2003; Teulon et al. unpublished data). They constitute a distinctive component of the world aphid fauna and New Zealand fauna, with species belonging to the Neophyllaphidinae, Taiwanaphidinae (primitive subtribes with Gondwanan distributions) and (mostly Aphidini). Recent molecular research places a group of native aphid species as central to the global evolution of the species-rich Aphidinae (von Dohlen & Teulon 2002) that includes many of the world pest species. New Zealand native species are not considered plant pests but those species in Aphis/Paradoxaphis and Euschiaphis genera are related to common pest species found in New Zealand. Aphid species native to New Zealand are generally monophagous (at least to plant genus) and are found on plant species bordering natural and productive systems.

Natural enemies.

The relatively narrow range of predators and parasitoids on aphids in NZ has been recognised for some time and as a result a number of predator (Coccinellidae, Chrysopidae, Hemerobiidae, Coniopterygidae, Chamaemyiidae and Syrphidae) and parasitoid (mostly Aphelinidae, , see below) species have been introduced into New Zealand for aphid pest management (Thomas 1989).

We are interested in the impact of introduced parasitoids on indigenous aphids and how this might inform future introductions of biological control agents into New Zealand. The New Zealand aphid/parasitoid system provides a useful model for assessing the non-target impact of natural enemies due to; (1) the relatively large numbers (in terms of species and total numbers) of introduced compared with native aphids found in New Zealand, (2) the number of parasitoid introductions that were made before the size of the native aphid fauna was known, (3) the close taxonomic relatedness of some native and introduced aphids in New Zealand, and (4) the type of host plants of New Zealand aphids that are often found on the border between natural and productive systems.

In this paper we review the known primary parasitoid fauna in New Zealand, including both deliberate and unintentional introductions of exotic parasitoids, and attempt to place these introductions in the context of their potential non-target impact on native New Zealand aphids.

PRIMARY PARASITOIDS: INTENTIONAL INTRODUCTIONS.

There have been seven or eight successful introductions, and a small number of unsuccessful intentional introductions of aphid primary parasitoids to New Zealand. These introductions are summarised in Cameron et al. (1989). One further introduction has taken place since then. Valentine and Walker (1991) and Anonymous (2007) provide a list of species established in New Zealand:

422 Proceedings of ISBCA 3 – P. G. Mason, D. R. Gillespie and C. Vincent Eds. (2008)

Aphelinus mali (Haldeman) (Aphelinidae) for the woolly apple aphid (Eriosoma lanigerum (Hausmann)) on apple. Sourced from several states in the USA and released in New Zealand from 1921 to 1924 after screening for secondary parasitoids. It became widespread in New Zealand by 1925 and is considered to be a highly successful biocontrol agent except where interrupted by pesticide applications (Walker 1989).

Aphelinus subflavens (Westwood) (Aphelinidae) for the oak aphid (Myzocallis annulatus (Hartig)) on oak and chestnuts. Introduced to New Zealand from Europe via Australia in about 1939, but subsequently found to be present in New Zealand at that time. The oak aphid is no longer considered a significant problem, suggesting that A. subflavens was a successful biocontrol agent (Walker 1989).

Aphidius eadyi Stary, Gonzales & Hall (Braconidae) for the pea aphid (Acyrthosiphon pisum (Harris)) on lucerne. Originated from Morocco via California and released into New Zealand from 1977 to 1981 with subsequent distribution throughout New Zealand. It became established in all lucerne areas by 1987, with parasitism rates of 30-40% associated with declining pea aphid populations (Cameron & Walker 1989).

Aphidius ervi Haliday (Braconidae) for the bluegreen lucerne aphid (Acyrthosiphon kondoi Shinji) and pea aphid (Acyrthosiphon pisum (Harris)). Strains from various countries (via California), Australia and UK were released into New Zealand from 1977 to 1981. A. eadyi and particularly A. ervi have contributed, along with a range of other natural enemies and resistant cultivars, to the control of the bluegreen lucerne and pea aphid (Cameron et al. 1989). It should also be noted that a specimen considered to be A. ervi was identified in New Zealand in 1963 from Aulacorthum solani on Histeropteris excelsa (a fern) (M. Carver pers. comm. to J Berry).

Trioxys complanatus Quilis (Braconidae) for the spotted alfalfa aphid ( trifollii (Monell)) on lucerne. Introduced into New Zealand from Australia from 1982 to 1985, established only in localised populations and has not been recovered since 1985. T. trifollii proved not to be a pest under New Zealand conditions (Walker & Cameron 1989).

Ephedrus plagiator (Nees ab Esenbeck) (Braconidae) was introduced from Japan via Australia for biocontrol of A. kondoi and A. pisum in 1977. Cameron et al. (1989) noted that field recoveries did not persist but this species is listed in the Checklist of New Zealand as being in New Zealand (Anonymous 2007) (M. Carver, pers. comm.).

Aphidius rhopalosiphi De Stefani Perez (Braconidae) for the rose-grain aphid (Metopolophium dirhodum (Walker) on cereals (esp. barley). Introduced from England and France and released in 1985 from 1987 with recoveries made from most cereal-growing areas in 1987 (Stufkens & Farrell 1989). This introduction was considered to be a success, with Grundy (1990) estimating that A. rhopalosiphi provided annual benefits of between NZ$0.3 and $5 million p.a. A. rhopalosiphi also parasitises another important cereal aphid pest found in New Zealand í Rhopalosiphum padi (L.).

423 Proceedings of ISBCA 3 – P. G. Mason, D. R. Gillespie and C. Vincent Eds. (2008)

Aphidius sonchi Marshall (Braconidae) for the sowthistle aphid ( (L.)) on blackcurrant. Introduced from Australia in 1994 and established throughout New Zealand, except for the southern South Island). Stufkens & Farrell (1995) suggest it is was likely to be present in New Zealand at the time of release. Significant levels of parasitism were recorded on its secondary host (sowthistle), but minimal levels were recorded on its primary host (blackcurrant).

Additionally, Aphidius smithi Sharma & Subba Rao, Ephedrus plagiator (Nees ab Esenbeck), and Praon barbatum Mackauer were introduced for biocontrol of A. kondoi and A. pisum in 1977, although there was no confirmation of their establishment (Cameron et al. 1989; Valentine & Walker 1991 Anonymous 2007).

It appears that the parasitoid impact on non-target aphid species was not examined until the introduction of A. rhopalosiphi in 1985 when this parasitoid species was only tested on other introduced aphid species (Table 1). Aphidius sonchi was tested on four of the known six New Zealand native species before its introduction in 1994 (Stufkens & Farrell 1994). All introductions were made before the extent of the indigenous aphid fauna became apparent in the late 1990s.

Table 1. Deliberate introductions of primary parasitoids for biocontrol of aphids – confirmed establishment in New Zealand. Host record data are from Carver 1984, 2000 for Australia (in Waterhouse & Sands 2001).

Parasitoid species Native aphids Introd Host aphid genera species (# tested) 1921 Eriosoma 0 (0) Aphelinus 1939 Tuberculoides 1 (0) subflavens Aphidius eadyi 1977 Acyrthosiphon 6 (0 a) Aphidius ervi Acyrthosiphon, Macrosiphon, 1977 6 (0) Metopolophium, Myzus Trioxys complanatus 1982 Therioaphis 6 (0) Aphidius Metopolophium, 1985 6 (0b) rhopalosiphi Rhopalosiphum Aphidius sonchi 1994 Hyperomyzus 6 (4c) 15+d

Information on the number of New Zealand native aphids and number of species tested from: aCottier 1953, bUnpublished files (Crop & Food Research), cStufkens & Farrell 1994, dTeulon et al. submitted.

PRIMARY PARASITOIDS: SELF-INTRODUCTIONS.

In addition to those primary parasitoids deliberately introduced into New Zealand for aphid biocontrol, a number of self-introduced species have established in New Zealand (Valentine & Walker 1991; Anonymous 2007) (Table 2).

424 Proceedings of ISBCA 3 – P. G. Mason, D. R. Gillespie and C. Vincent Eds. (2008)

Examples include:

Diaeretiella rapae (M’Intosh) (Braconidae) has been recorded in New Zealand since at least 1930 (Gourlay 1930) and is a parasitoid of Brevicoryne brassica. Its effectiveness as a biocontrol agent has been severely undermined by a number of hyperparasitoids (Cottier 1953; Thomas 1989).

Aphidius salicis Haliday (Braconidae) is a parasitoid of the carrot aphid (Cavariella aegopodii (Scopoli)), which has been recorded in New Zealand since at least 1962 (Carver & Stary 1974). While carrot aphid is a serious pest of carrots in New Zealand (Lowe 1971), there is little information on the impact of the parasitoid. However, in Australia the intentional introduction of A. salicis in 1962 and a change in the predominant carrot cultivar grown significantly reduced the pest status of the carrot aphid (Waterhouse & Sands 2001).

Aphidius colemani Viereck (Braconidae) is not recorded in Valentine & Walker (1991), but is found in Anonymous (2007). Unpublished records suggest it has been in New Zealand since at least 1982. It is an effective biological control agent for a range of aphid species in greenhouse crops and is commercially available in New Zealand (www.bioforce.net.nz, www.zonda.net.nz).

Praon necans Mackauer (Braconidae) is a central and eastern European species that attacks Rhopalosiphum and Schizaphis species. This species, or one very similar to it, has been found parasitising a New Zealand native Euschizaphis species (Mackauer pers. comm.), but has not been recorded from any introduced species in New Zealand.

There are also a number of unpublished records of the following exotic aphid parasitoid species in New Zealand: Ephedrus persicae, Trioxys auctus, Trioxys cirsii, Trioxys compressicornis, Trioxys pallidus and Trioxys tenuicaudus.

PRIMARY PARASITOIDS: INDIGENOUS APHIDS.

Field surveys.

Since about 1998 we have made observations on and collected parasitoids, including primary parasitoids, of native New Zealand aphids. Approximately 400 parasitoids have been collected from a range of native aphid species (Table 3). Most were collected in the South Island of New Zealand, where most native aphid populations have been found. We have concentrated our efforts on the New Zealand native Aphis/Paradoxaphis/Casimira group, which is closely related to many pest species.

DNA diagnostics.

Limited knowledge about the New Zealand aphid parasitoid fauna and the lack of available aphid parasitoid taxonomic expertise has hampered efforts to measure the non-target impacts of introduced biocontrol agents.

425 Proceedings of ISBCA 3 – P. G. Mason, D. R. Gillespie and C. Vincent Eds. (2008)

Table 2. Self-introductions of primary parasitoids – confirmed establishment in New Zealand. Host record data is from Carver 1984, 2000 for Australia (in Waterhouse & Sands 2001).

Parasitoid species Host aphid genera NZ reference Braconidae Aphis, Myzus, Rhopalosiphum, Aphidius colemani Anonymous 2007 Toxoptera Valentine & Walker Aphidius pelargonii Acyrthosiphona 1991 Aphidius salicis Cavariella Carver & Stary 1974 Valentine & Walker Aphidius similis Myzus, Rhopalosiphum 1991 Brevicoryne, Myzus, Gourlay 1930, Cottier Diaeretiella rapae Rhopalosiphum 1953 Lysiphlebus Aphis, Rhopalosiphum, Anonymous 2007 testaceipes Toxoptera Praon nr. necans [Euschizaphis]b Mackauer pers. comm. Aphelinidae Valentine & Walker Acyrthosiphon 1991 Valentine & Walker Therioaphis 1991 Aphis, Macrosiphum, Myzus, Valentine & Walker Aphelinus gossypii Toxoptera 1991 Valentine & Walker Aphelinus humulis Aphis 1991

aNew Zealand host record from Valentine & Walker (1991). bNew Zealand host record.

Our research group has been trialling a DNA sequence-based diagnostic technique for identifying aphid parasitoids to species level. A reference data set consisting of mitochondrial 16SrRNA sequences has been assembled. These data comprise available sequences from braconid and aphelinid taxa that appear in the New Zealand Collection checklist of New Zealand hymenoptera (Anonymous 2007). New Zealand taxa not represented in public domain sequence databases are represented by sequences from congeners.

Wasp-specific primers (Jones et al. 2005) were used to detect the presence of a parasitic in mummies collected from the field (see above). Amplified fragments were gel purified and sequenced. Following sequence editing, ClustalX (Thompson et al. 1997) was used to align mummy 16S sequences with those in the parasitoid reference set. PAUP* (Swofford 2000) was used to calculate a genetic distance matrix from which a neighbour-joining topology was generated.

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Parasitoids on native aphids.

Results to date indicate that DNA sequence data is a useful tool for cataloguing the diversity of the New Zealand native aphid parasitoid fauna as a range of braconid and aphelinid species have been identified from our field surveys (Table 3). Some species have yet to be fully characterised and may be New Zealand native parasitoids new to science and other species need to be verified using morphological characters. The overall impact of any of these parasitoids on native aphid populations is unknown at this time . Introduced parasitoids have now been implicated attacking three New Zealand native aphid species: (1) Aphidius nr ervi on Aphis cottieri (Carver 2000), (2) Several Aphidius species on Aphis healyi, and (3) Praon nr. necans on a Euschizaphis species (M. Mackauer, pers. comm.). Given the taxonomic relatedness of several native aphid species to introduced aphid pest species and the polyphagous nature of several aphid parasitoids it is not unexpected that native aphids are attacked by introduced parasitoids. A. ervi is selectively polyphagous on the but is also know to attack Aphis species (Marsh 1977). Eushizaphis is closely related to Rhopalosiphum and Schizaphis which are known hosts for Praon necans (M. Mackauer, pers. comm.).

Table 3. Primary aphid parasitoids attacking selected indigenous aphids in New Zealand.

Native aphid species Primary parasitoid Reference/comments Aphis coprosmae Aphidius sp. No aphids or parasitoids Aphelinus sp. collected since 1997 Aphis cottieri Aphidius nr. ervi Carver 2000 Aphidius sp.a Aphis healyi Aphidius sp.a,b Aphidius sp.a Aphis nelsonensis None recorded Aphid colonies not seen since 1965 Aphis sp. (on Olearia) Aphelinus sp.a Aphis sp. (on Hebe) No parasitoids Aphid colonies recently recorded to date discovered 2008 Paradoxaphis aristoteliae Not yet subjected to DNA analysis Paradoxaphis plagianthi Not yet subjected to DNA analysis Euschizaphis spp. Praon nr. necans Mackauer pers. comm. (Dracophyllum) Euschizaphis sp. (Aciphylla) Aphidius sp.a Lysiphlebus sp.a Neophyllaphis totarae New genus/species Mackauer pers. comm. Neophyllaphis sp. (on P. nivalis) Sensoriaphis nothofagi None recorded

aParasitoid species confirmed by DNA sequence data. bPutative introduced species.

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CONCLUSIONS.

A range of primary aphid parasitoid species (a mixture of intentionally introduced species, self-introduced species and probably indigenous species) are recorded from introduced and indigenous aphids in New Zealand. Both deliberately and unintentionally introduced parasitoids species make important contributions to the biological control of pest aphid species. Lack of knowledge of New Zealand native aphids and aphid parasitoids, including their , is a severe impediment to understanding the impact of introduced parasitoids on native aphids and how this information might inform future introductions of biological control agents into New Zealand.

ACKNOWLEDGEMENTS.

We thank Manfred Mackauer for assistance in parasitoid identification. This work was partially funded by New Zealand’s Foundation for Research, Science and Technology through the Better Border Biosecurity (B3) Programme (www.b3nz.org).

REFERENCES.

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