recent non-native invertebrate plant pest establishments in great britain: origins, pathways, and...

© 2007 The Authors

Journal compilation © 2007 The Royal Entomological Society

Agricultural and Forest Entomology (2007), 9, 307–326

Introduction

The unintentional human-assisted movement of invertebrate plant pests between countries has been well documented ( Elton, 1958; Sailer, 1983; Baker & Aitkenhead, 1986; Brockerhoff et al. , 2006 ). This awareness is reflected in a well-established international agreement for preventing the movement of plant pests, the International Plant Protection Convention (IPPC; FAO, 1997 ). Although there have been considerable successes in limiting the movement of high risk (i.e. quarantine) species, Spain ( Pérez Moreno, 1999 ), France ( Martinez & Malausa, 2000; Streito & Martinez, 2005 ), Italy ( Pelizzari et al. , 2005 ) and Serbia and Montenegro

( Glavendeki ć et al. , 2005 ) have reported recent increases in the establishment rates of plant pests. Although it is difficult to determine the extent to which such increases are due to changes in sampling effort ( Pelizzari & Dalla Monta, 1997 ), the ongoing flow of non-native plant pests between countries has also been recorded in the U.S.A. ( OTA, 1993 ), Japan ( Kiritani & Yamamura, 2003 ) and Australia ( Maynard et al. , 2004 ). This is considered largely to be a consequence of the globalization of trade. The volume of international trade and travel is so great, and the modes of entry are so varied, that not all consignments or routes of entry can be screened ( Levine & D’Antonio, 2003; Work et al. , 2005 ). Rather, the resources of plant protection organizations are focussed on priority pathways, whereas new threats may arise from novel markets or commodities.

According to the ‘tens rule’ ( Williamson & Fitter, 1996 ), ap-proximately 10% of non-native species that arrive eventually

Recent non-native invertebrate plant pest establishments in Great Britain : origins, pathways, and trends

Richard M. Smith , Richard H. A. Baker , Chris P. Malumphy , Sue Hockland , Roger P. Hammon , Joe C. Ostojá-Starzewski and Dominique W. Collins Central Science Laboratory, Sand Hutton, York YO41 1LZ, U.K.

Abstract 1 An appraisal of non-native invertebrate plant pest establishments in Great Britain, between 1970 and 2004, was carried out to improve our understanding of current invasion processes by non-native plant pests, and to assist national strategies in managing the risks they pose.

2 A total of 164 establishments, comprising 50 natural colonists and 114 human-assisted introductions, were recorded across 13 major taxonomic groups.

3 The mean rate of establishment was 22.1 species per 5-year period: 19.1 and 3.0 species outside and inside protected cultivation, respectively. Despite the contin-uing rapid growth in international trade and a general perception that rates of pest invasions are accelerating, no significant temporal trends in the rate of estab-lishments in Great Britain were detected, either for natural colonists or human-assisted introductions, or for pests of plants grown indoors or outside.

4 The plant trade, particularly in ornamental plants, accounted for nearly 90% of human-assisted introductions; apiculture, biological control, timber imports, transport stowaways and intentional releases each contributed less than 5%. Only eight (4.9%) of the establishments could be considered as having no direct poten-tial economic impact because all other species have been recorded as feeding on cultivated plants. A greater proportion of establishments by both natural colonists and human-assisted introductions occurred on non-native, woody plants.

5 The present study confirms previous work in other European countries that highlight the predominant role of the ornamental plant trade in introducing new plant pests to the European continent, mainly from Asia and North America.

Keywords Flatworms , insects , invasive alien species , mites , nematodes , plant health .

Correspondence: Richard M. Smith, 12 Chestnut Grove, Acomb, York YO26 5LE, U.K. Tel: + 44 (0) 1904 793679; e-mail: [email protected]

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© 2007 The Authors

Journal compilation © 2007 The Royal Entomological Society, Agricultural and Forest Entomology, 9, 307–326

establish and, of these, a similar percentage will cause serious economic or environmental impacts. Thus, basic knowledge of the identity, number, origin, and entry pathway of species that establish is essential for assessing threats from non-native spe-cies. Such a need is addressed by the IPPC, which requires na-tion states to maintain adequate information on pest status (Article VII, paragraph 2j; FAO, 1997 ); and the Guiding Principles of the Convention on Biological Diversity (CBD) stipulate an adequate knowledge base for non-native species (Sixth Conference of the Parties, Decision VI/23; CBD, 2002 ). Consequently, for example, the U.K. government’s review of national policy on non-native species made monitoring and surveillance a key recommendation for limiting impacts ( Defra, 2003 ).

There are no up-to-date species lists of invertebrates inju-rious to plants recently established in Great Britain. A list of introduced organisms ( Brown, 1986 ) and records of non-native invertebrate herbivores held in the Phytophagous Insect Data Bank ( Ward, 1988 ) are inconsistent, contain nu-merous omissions, or are out of date. Recent audits of non-native species in Scotland ( Welch et al. , 2001 ) and England, with contributions from the present study ( Hill et al. , 2005 ), excluded species found only in protected environments (e.g. glasshouses, and non-native pests that had spread by natural colonization). Smith et al. (2005) provided a preliminary analysis of the data reported here. The present study updates and extends this analysis, providing a complete list of the in-vertebrate pest establishments recently recorded in Great Britain.

The present study provides the first up-to-date evidence of the non-native plant pests that have established in Great Britain. Such evidence can contribute to assessments of the relative risks of different introduction pathways and com-modities in a developed, Western European state. Great Britain is a particularly appropriate region in which to com-pare the relative effects of natural colonization and human-assisted introduction; its island status allows a clearer delineation between the two modes of arrival compared with continental countries. Furthermore, a strong tradition of nat-ural history study means that expert amateurs contribute a wealth of information on new discoveries, spanning a wide range of taxonomic groups ( BRC, 2006 ).

The present study had three specific aims. First, to create a database of non-native invertebrate plant pests establishing in Great Britain from 1970 to 2004 to identify any trends over time. Both intentional and unintentional introductions were included in our study, as were natural colonizers be-cause, unlike the CBD Guidelines on Invasive Alien Species ( CBD, 2002 ), IPPC standards on phytosanitary measures are not restricted to species that have entered an area through man’s activities ( Baker et al ., 2005 ). Also, distinguishing human-assisted introduction from natural colonization is critical for devising appropriate plant health responses. Trade pathways can be managed through phytosanitary reg-ulation, but preventing establishments after natural dispersal depends primarily on eradication measures. The present study differs from previous inventories of non-native species in Great Britain by including pests established in protected environments. A proportion of such species have been able

to spread outside, both in Great Britain and elsewhere in Europe (Pellizzari & Dalla Montà, 1997; Kenis, 2005 ), and this may be enhanced as climate change provides hotter summers for development and warmer winters for survival ( Baker et al ., 2003 ).

Second, to analyse trends in the numbers of plant pests es-tablishing from different continents and pathways. By track-ing changes in the numbers of pests from source areas, it is possible to quantify threats from emerging or declining markets ( Thomas, 2000; Levine & D’Antonio, 2003; Normile, 2004 ), and hence focus limited resources. Similarly, monitor-ing establishments in relation to pathways enables the threats of different modes of entry to be addressed.

Third, to identify those non-native plant pests that threaten the economy, by feeding on cultivated plants, or the environ-ment by attacking uncultivated plants. The potential impacts of pests were explored through an analysis of their host spe-cificity, of the cultivated status of hosts, and of the type of environments where the hosts occurred. Host specificity was used to assess potential impact because information on host damage (e.g. mortality and reduction in yield and quality) is unavailable for the majority of species, as is often the case for all but the most high-profile pests ( Moran, 1983 ).

Materials and methods

Data sources

Published and unpublished information on non-native inver-tebrate plant pests was extracted from the professional and amateur literature (particularly checklists of the British fauna) and Central Science Laboratory (CSL, Defra) files. CSL opens official case files on pest species if they have been intercepted at ports of entry, or if outbreaks on plants have been reported. Stored product pests and those attacking cut timber were omitted. In addition to herbivores, the IPPC also defines species that harm organisms beneficial to plants as pests ( FAO, 2002 ). Therefore, where a risk or evidence of negative impact had been identified by a national plant pro-tection organization, predators and parasites of pollinators, earthworms and beneficial natural enemies were also included.

The subset of the database used here consisted of plant pests, newly-established subsequent to 1970, that have main-tained self-sustaining populations in indoor (e.g. glasshouses) or outdoor environments (e.g. gardens, farmland, and other cultivated and uncultivated habitats). The data for this period are more robust than earlier years because their sources are generally more accessible in on-line bibliographic databases. Species currently under official control (i.e. eradication or containment measures, reintroductions of extinct natives), migrants and species with only transient populations (casu-als) were excluded. Because the herbivores of higher plants are relatively well known compared with many other inverte-brate taxa in Great Britain ( Hill et al. , 2005 ), expert reviews are generally able to distinguish new, non-native species from overlooked natives ( Kirby et al. , 2001 ). New species of un-certain status, Furcipus rectirostris ( Read, 1981 ), Ixapion

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variegatum ( Foster et al. , 2001 ), Pristiphora leucopus ( Grearson, 2006 ) and taxonomic revisions were excluded from the analyses.

The dataset included plant pests recorded up to 2004. This threshold was chosen partly for convenience in analysing es-tablishments over 5-year periods, and partly because the lag in reporting a first detection of a species is approximately 1 – 2 years (median 2 years, modal class 1 year, for the period 1970 – 2004). Therefore, the value for the number of estab-lishments in 2000 – 2004 is unlikely to be a significant under-estimate of the true value. Although the dates of detection are conventionally used as the date of establishment, the true es-tablishment date may have occurred some time previously, with the length of the detection lag being dependent on re-cording effort.

Some definitions of newly-arrived species distinguish es-tablished from transient populations by their persistence for more than 4 years ( Hill et al. , 2005 ). Species that arrived af-ter 2001 were included if the available evidence indicated that they were likely to have established. Evidence from the Lepidoptera suggests that most new species established long-term populations (86 out of 89 establishments in the period 1900 – 1999; Parsons, 2003 ).

Mode of entry

New species established in Great Britain were categorized as having entered and colonized by natural spread (i.e. by active or passive aerial dispersal) or with human assistance (either in-tentional or unintentional). The categorization followed the ev-idence provided in published sources and, where possible, multiple sources were consulted to arrive at the assessment. In some cases, the opinion of a national expert was sought. For some species, a definite statement on the mode of entry was unavailable; it is also probable that some species entered Great Britain by more than one mode of entry. In these cases, the ele-ment of uncertainty was noted (see Appendix 1a and b ), and the likely principal means of establishment was adopted. Species that may have colonized Britain naturally from a re-gion where they had been introduced by humans (e.g. harle-quin ladybird, Harmonia axyridis ; Majerus et al. , 2006 ) were considered as having entered through human assistance, and termed ‘indirect introductions’. Information on the pathway of entry (e.g. type of commodity in trade) was gathered at this stage.

Native range

We identified the continent where each pest is native. Those species with worldwide distributions, possibly resulting from undocumented introductions, were recorded as ‘cosmopoli-tan’. The source of entry into the U.K. is often very difficult to establish (although the origin may be the same as the source) because non-native introductions do not always oc-cur directly between two countries, but via other states. Information on the native range may nevertheless identify re-gions that pose a high risk as a source of non-native pests (e.g. by providing evidence of particular trade pathways).

Regions of Britain where pests were first detected

The distributions of first records of established species were collated in accordance with the regional definitions and vice-counties in Hill et al. (2005) . English regions were groupings of Watsonian vice-counties ( The Ray Society, 1969 ), except for ‘London’, which comprised Middlesex and north Surrey. In the present study, Middlesex was incorporated into the re-gion ‘south east’ to maintain coherency with vice-county boundaries. The single first establishment in Scotland was recorded as ‘Scotland’.

Cultivated and noncultivated hosts

The potential impacts of pests were assessed according to whether their hosts are cultivated (e.g. agriculture/horticulture and forestry, noncultivated or both). A pest was categorized as having potential economic and environmental impacts in two situations: if it had a single host that was both cultivated and occurring naturally (e.g. Tinagma balteolella on Echium vulgare ; Agassiz, 1976 ), or if it had multiple hosts that included cul-tivated and wild plants (e.g. Gelechia senticetella on Juniperaceae and Cupressaceae; Emmet & Langmaid, 2002 ).

Host specificity

Host specificity is another measure of a pest’s potential threat. Although monophagous species can be very damaging, the wider the host range, the greater the number of species that could suffer economic or environmental damage. Host range was determined from the literature and, as such, was poten-tially wider than has been recorded in Great Britain itself. This estimate was used because it indicates the capacity of a pest to exploit hosts, even if the pest is initially restricted in its host range (e.g. Phyllonorycter leucographella spread to other ro-saceous hosts after its invasion on Pyracantha spp.; Palmer, 1999 ). Similarly, Epiphyas postvittana , a serious apple pest in Australia and South Africa, has only been found on ornamen-tals in Great Britain after its invasion in 1936 ( Alford, 1984 ), although it will feed on apple in captivity ( Porter, 2001 ).

When describing host range, herbivores are conventionally distinguished between those that are monophagous (feeding on one host taxon), oligophagous (a few host taxa), or poly-phagous (many host taxa). Because these terms are relative ( Gullan & Cranston, 2005 ), we adopted the following terms to allow comparisons with other studies of the British fauna ( Ward & Spalding, 1993 ): feeding on species of one host genus (monophagous), one to three plant families (oligopha-gous) or more than three plant families (polyphagous). Typically, most oligophagous herbivores feed on a range of genera within a single plant family. Host specificity was compared between pest taxa to determine whether certain taxa threaten a wider range of hosts.

Host growth form

The relationship between the growth forms of hosts (on which pests first established) and their alien or native status was explored. Ornamental plants, dominated by alien


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species, are a major source of plant pest introductions (Pellizzari & Dalla Monta, 1997). Such plants include a dis-proportionate number of woody species (trees and shrubs) compared with annuals, biennials and perennials, both in gar-den environments ( Smith et al. , 2006 ) and the British alien flora ( Crawley et al. , 1996 ). Thus, it might be expected that a disproportionate number of British establishments are associ-ated with woody aliens. This was tested by comparing the proportions of alien and native hosts in each category of growth form against the proportions of alien and native hosts across the entire dataset.

Growth form was examined further by comparing the ra-tios of different growth forms of native and non-native unculti vated plants in the British flora (the ratio of annuals/biennials : perennials : woody plants is 23 : 58 : 19 for al-iens, and 24 : 67 : 9 for natives; Crawley et al. , 1996 ) with those used as hosts by natural colonists. Assuming random selection of the pool of potential hosts by a pool of potential natural colonizers, no departures would be expected from the ratios of growth forms in available hosts in uncultivated hab-itats. The analysis was conducted only on natural colonists because most human-assisted introductions were, by defini-tion, linked to a host plant. Knowledge of the relative fre-quencies of growth forms moved in trade would have been required, but such data are unavailable.

Statistical analysis

G -tests of independence were used to identify statistically significant associations between categorical variables ( Sokal & Rohlf, 1995 ). Analyses of particular pairs of variables were restricted to those classes possessing cell totals ³ 5, and the significance of G was tested against the chi-square distribu-tion ( Rohlf & Sokal, 1995 ). Linear regression was used to explore trends in established numbers over time. There was no evidence for the need to include autoregressive terms in models (S. Pietravalle, personal communication).

Results

The taxonomy and establishment status of pests

A total of 164 invertebrate plant pests were recorded from 13 major taxonomic groups ( Table 1 ). They were dominated by the Homoptera (33.9%) and Lepidoptera (25.5%), and just four insect orders together contributed more than 80% of all establishments. Establishments were divided into two modes of entry, either natural colonists (30.3%) or introductions as-sociated with man (69.7%). Mode of entry was not random with respect to taxonomic group ( G = 82.358, d.f. = 6, P < 0.001): human-assisted introductions consisted disproportion-ately of Homoptera and Coleoptera, whereas natural colonists were mainly composed of Lepidoptera and Heteroptera.

Trends in establishments over time

The number of establishments fluctuated around a mean ( ± SE) of 22.1 ± 2.06 per 5-year period ( Fig. 1 ), ranging between a

minimum of 15 (1985 – 1589) and maximum of 31 (1995 – 1999). Establishments were below the long-term average in three out of four of the 5-year periods before 1990, whereas all were above the long-term average after 1990. There were no significant trends in the number of species establishing over the period 1970 – 2004, either for total numbers ( F 1,5 = 1.05, P = 0.35), or for the separate modes of entry: human-assisted introductions (mean 15.0 ± 1.40; F 1,5 = 2.82, P = 0.15), and natural colonists (mean 7.14 ± 1.16; F 1,5 = 0.0, P = 0.97).

There were relatively few establishments in protected en-vironments (e.g. in glasshouses) compared with outside (17.1%); the mean number ( ± SE) per 5-year period was 3.0 ± 0.87 for species establishing indoors or both indoors and out-side, and 19.1 ± 2.10 for species only establishing outside. As for modes of entry, there were no significant trends over time for the numbers of species establishing either indoors (including both indoors and outside: F 1,5 = 1.40, P = 0.29) or outside ( F 1,5 = 0.24, P = 0.64).

Table 1 Mode of entry in relation to the taxonomic status of plant pests established in Great Britain

Taxon Human-assisted introductions

Natural colonists Total %

Homoptera 53 3 56 34.1 Lepidoptera 13 28 41 25.0 Coleoptera 17 2 19 11.6 Heteroptera 1 14 15 9.1 Diptera 8 0 8 4.9 Acari 6 2 8 4.9 Hymenoptera 4 1 5 3.0 Nematoda 4 0 4 2.4 Phasmatodea 2 0 2 1.2 Platyhelminthes 2 0 2 1.2 Thysanoptera 2 0 2 1.2 Blattodea 1 0 1 0.6 Mollusca 1 0 1 0.6 Total 114 50 164 100.0

01970-4 1975-9 1980-4 1985-9 1990-4 1995-9 2000-4

5

10

15

20

25

30

35

No.

of s

peci

es e

stab

lishe

d

Indirect introductionsDirect introductions

Colonists35 year mean

Figure 1 Trends in the numbers of plant pests establishing in Great Britain (1970 – 2004), in relation to establishment status ( n = 155, missing data, n = 9). ‘Indirect introductions’ are species establishing by natural spread, but initially introduced elsewhere.


© 2007 The Authors


Geographic origins of pests

Nearly half of all established species (49.7%) originated in mainland Europe ( Table 2 ), and this region was the source of virtually all the natural colonists. Other major contributing regions for human-assisted introductions were Asia and North America (each with 13.3%), whereas the origins of a substan-tial proportion of species were unknown (10.9%). No trends were apparent for changes in the source of establishments over the period 1970 – 2004 ( Table 2 , for regions providing more than ten species: Europe, F 1,5 = 0.974, P = 0.37; Asia, F 1,5 = 2.47, P = 0.18; North America, F 1,5 = 0.455, P = 0.53).

Regions of Britain where pests were first detected

Most establishments ( n = 162; two in no single region) were first detected in the south-east (53.1%) and east (14.8%) of England, followed by the south-west (8.6%), north-west (5.6%), west-midlands (1.2%), Yorkshire and the Humber (0.6%), north-east (0.6%) and Scotland (0.6%); the locations for a substantial number were unknown (14.8%). Only in the north-west did the ratio of natural colonists to human-assisted introductions depart from random expectation, based on pro-portions across all regions ( G = 6.708, d.f. = 1, P < 0.01): no natural colonists were first detected in the north-west. Indeed, natural colonists were confined to the southern portion of England (i.e. south-east, east and south-west). Concerning hu-man-assisted introductions, 47.3% occurred in the south-east, 11.6% in the east, 7.1% in the south-west and 8.0% in the north-west, whereas only 4.5% were reported from the other regions combined. Regional totals also hid finer scale patterns discernable at the Watsonian vice-county level. For example, in the south-east, East Kent held 42.4% of all natural colo-nists in the region ( n = 14), whereas Surrey alone contained 45.3% of all the human-assisted introductions ( n = 24).

Entry pathways

A substantial majority of the pests introduced by human activity were associated with the movement of plants (89.0%).

Other pathways made relatively insignificant numerical con-tributions: apiculture (1.8%), biocontrol (0.9%), intentional releases (1.8%), timber imports (2.8%) and transport stowa-ways (3.7%). Human-assisted introductions with ornamental plantings outside (e.g. gardens, parks, amenity, landscaping; 58.8%) far exceeded those introductions associated with for-estry (4.38%), agriculture/horticulture (1.75%), or with api-culture (1.75%) ( Table 3 ). Only seven out of 24 establishments under cover were linked with horticultural crops. Thus, taken together, all ornamental plants, both outside and under cover, accounted for 73.7% of all human-assisted introductions with plants. By contrast, natural colonists mainly established in the wider countryside ( Table 3 ), with small minorities in or-namental (e.g. Thera cupressata on Cupressus spp.; Waring, 1998 ) and forestry environments (e.g. Dioryctria schuetzeella on Picea abies ; Parsons, 2003 ).

The potential impact of pests based on cultivation status of hosts

When assessing potential economic and environmental threats of plant pests from the types of hosts affected, pests of cultivated and uncultivated plants would be expected to pose mainly economic and environmental threats, respec-tively. The distribution of threats to the different groups of plants was indeed nonrandom ( G = 21.14, d.f. = 2, P < 0.001). Predictably, pests under cover pose only economic threats ( Table 3 ). However, although most pests in ornamen-tal situations (e.g. parks, gardens and landscaping) also pose only economic threats (56.7%), a substantial minority have potential impacts on both the economy and environment (43.3%). Similarly, whereas few pests of hosts in the wider countryside pose only economic threats (13.5%), relatively few pose only environmental threats (15.4%), and most have a combined threat (71.2%).

Host specificity and taxonomic group of plant pest

The degree of host specificity was not random with respect to invertebrate taxon ( Table 4 ; for taxa with more than five

Table 2 Geographic origins of plant pests established in Great Britain, in relation to mode of entry ( n = 164) and period of first detection (1970 – 2004; n = 155, nine species lack precise dates of detection)

Origin Int’n Col’t 1970 – 1974 1975 – 1979 1980 – 1984 1985 – 1989 1990 – 1994 1995 – 1999 2000 – 2004 Total (years)

Europe 39 43 14 8 10 7 14 16 13 82 North America 22 0 7 2 3 1 2 3 4 22 Asia 22 0 2 1 1 1 3 8 3 19 Unknown 13 5 1 5 0 2 2 3 1 14 Australasia 9 0 0 2 2 1 3 1 0 9 Cosmopolitan 4 1 0 0 1 0 0 0 3 4 Americas (tropical)

3 0 0 0 0 2 0 0 0 2

Africa 1 1 1 0 1 0 0 0 0 2 South America 1 0 0 0 0 1 0 0 0 1 114 50 25 18 18 15 24 31 24

‘Europe’ includes Mediterranean species except those of exclusively North African origin. ‘Unknown’ includes species whose native origin is stated as such in the literature, or whose origin could not be verified from the literature for the present study. Int’n, human-assisted introduction; Col’t, natural colonist.


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establishments, G = 21.59, d.f. = 12, P < 0.05). The Homoptera contained disproportionately more oligophagous and polyphagous species compared with other taxa ( G = 7.792, d.f. = 2, P < 0.025). Although the Lepidoptera ap-peared to possess more monophagous species than expected, the proportion was marginally not significant ( G = 5.704, d.f. = 2, P < 0.1).

Establishments in relation to plant host status and growth form

The establishment of pests on different growth forms of host plant (i.e. annual or biennial, perennial or woody) also depended on the alien or native status of the host ( G = 39.54, d.f. = 2, P < 0.001): native hosts were more likely to be annuals/biennials or perennials, whereas alien hosts dis-proportionately comprised woody species (trees or shrubs, Fig. 2A ).

Concerning natural colonists, establishments on alien and native hosts were as expected from the ratios of aliens and natives in the British flora (establishments: on aliens 39.6%, on natives 60.4%; flora: aliens 43.6%, natives 56.4%; G =

0.312, d.f. = 2, not significant). However, among the differ-ent growth forms of alien plants, trees and shrubs were more likely to be colonized than either annuals/biennials or peren-nials ( G = 49.570, d.f. = 2, P < 0.001; Fig. 2B ). Among na-tives, there was no departure from the proportions found in the wild ( G = 0.906, d.f. = 2, not significant).

Discussion

The analyses of British invertebrate plant pest establishments highlight: (i) the lack of a discernable increase or decrease in establishments over the period 1970 – 2004; (ii) the substan-tial contribution by natural colonists to establishments; (iii) the broad potential threat of pests to both cultivated and un-cultivated plants; and (iv) the disproportionate number of plant pests that have established on woody plants, compared with other growth forms, even among natural colonizers. In addition, the present study has shown that: relatively few in-sect orders dominate establishments; Europe, North America and Asia are the major sources of new pests, but with no sig-nificant changes over time in the rates of introduction from

Table 3 Environments associated with establishments of plant pests, in relation to the mode of entry and the potential impacts resulting from damage to principal hosts

Environment Int’n Col’t Econ’c Env’l Both n/a Unknown Total (impacts)

Ornamental 65 6 38 0 29 1 3 71 Uncultivated habitats 14 41 7 8 37 1 2 55 Under cover 24 0 23 0 0 1 0 24 Forestry 5 3 5 0 3 0 0 8 Agriculture/horticulture 2 0 1 0 1 0 0 2 Apiculture 2 0 2 0 0 0 0 2 Unknown 2 0 0 0 1 0 1 2 Total 114 50 76 8 71 3 6 164

‘Ornamental’ refers to establishments on hosts planted outside for aesthetic reasons, ‘Agriculture/Horticulture’ refers to hosts planted as crops outside, either commercially or for personal consumption. Int’n, human-assisted introduction; Col’t, natural colonist; Econ’c, economic; Env’l, environmental; ‘n/a’, refers to parasites or predators.

Table 4 Degrees of host selectivity in established plant pests

Taxon Monophagous Oligophagous Polyphagous Unknown n/a Total

Homoptera 17 26 9 4 0 56 Lepidoptera 29 11 1 0 0 41 Coleoptera 7 5 3 3 1 19 Heteroptera 6 6 1 1 1 15 Acari 4 2 1 0 1 8 Diptera 6 1 0 0 1 8 Hymenoptera 4 1 0 0 0 5 Nematoda 2 0 0 2 0 4 Phasmatodea 0 0 0 2 0 2 Platyhelminthes – – – 0 2 2 Thysanoptera 0 0 2 0 0 2 Blattodea 0 0 0 0 1 1 Mollusca 0 0 1 0 0 1 75 52 18 12 7 164

Monophagous, restricted to one plant genus; oligophagous, restricted to £ 3 plant families; polyphagous, occurring on >3 plant families; n/a, omnivorous, carnivorous or parasitic species.


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these regions; and the ornamental plant trade acts as the most significant source of new pests.

Concerning data quality in the present analysis, Great Britain lacks published checklists for certain relatively poorly studied groups of plant pests, such as the Nematoda and Acari (with the exception of gall-formers; Redfern et al. , 2002 ). There is also a paucity of up-to-date distributional data for these groups because expert amateurs tend not to cover these taxa. Therefore, the numerical contribution of these groups to the analysis needs to be interpreted cautiously.

In general, as for any country ( Kenis, 2005 ), it is exceed-ingly difficult to compile an accurate database containing all the non-native plant pests that have established within a na-tion’s territory. Even if all the original sources of reports can be read to ensure that the new record is not a taxonomic revi-sion, or the new finding of a species that is likely to be na-tive, some records will remain unclear. We believe that, by consulting multiple sources for as many species as possible (including national experts in some cases), the number of misclassifications was minimized. Undoubtedly, future work will be able to clarify the status of certain species At this point, it is vital that any uncertainties are open to scrutiny,

and that advances in knowledge are linked to the same base-line information.

Trends in establishments

Despite marked fluctuations between periods, no significant trends in British establishment rates were detected in the num-bers of plant pests establishing in 1970 – 2004 for natural colo-nizers, human-assisted introductions, or for species under cover or outside. The mean establishment rate remained at more than four species per year, with 50 natural colonists and 114 human-assisted introductions arriving in Great Britain during the 35-year period studied. Direct comparisons with other European countries are hampered by the scope of stud-ies and definitions used, although the British data for intro-ductions are of a very similar order of magnitude to the 132 establishments recorded in Italy ( Pelizzari et al. , 2005 ) and the 94 in France (Martinez & Malausa, 2000; Streito & Martinez, 2005 ). If approximately 10% of establishments be-come significant pests under the ‘Tens rule’ ( Williamson & Fitter, 1996 ), it would be predicted that approximately 16 of the British species will cause substantial economic or environ-mental impacts. Smith et al. (2005) identified approximately one-half of the post-1970 establishments in Great Britain as being of sufficient plant health significance to merit the col-lection of information; however, there are insufficient data and no recognized thresholds for determining whether impacts have been substantial.

Although strictly comparable data are unavailable, rela-tively few establishments occurred in protected environments in Great Britain (17.1%) and in Italy (19.7%, 1945 – 2004; Pelizzari et al. , 2005 ), compared with the Netherlands, where at least 30 out of 72 species (41.7%) have established in pro-tected environments subsequent to 1900 ( Van Lenteren et al. , 1987 ). Although the relatively low rate of British establish-ments under cover, generally commercial glasshouses and botanic gardens, may reassure these sectors, approximately 20% of species establishing under cover later spread outside, and so the risks posed by these species may not be confined to protected environments. This phenomenon has been ob-served particularly in countries with warmer climates ( Kenis, 2005; Pelizzari et al. , 2005 ).

Both Italy and France have reported increased rates of es-tablishment in recent years, but it is unclear what role sam-pling intensity has played in these changes (see below). Florida is a well known recipient of new pests, but even here an accelerating establishment rate has not been found during the last 15 years ( Loope & Howarth, 2003 ); a result corrobo-rated by evidence on the pests found on vegetables in the U.S.A. ( Carpinera, 2002 ). It is surprising not to have found accelerating establishment rates in Great Britain because there have been dramatic increases in the volume of plants and plant products imported to the U.K. between 1970 and 2004. For example, fresh fruit and vegetable imports have more than doubled from 2 571 345 tonnes in 1970 to 5 634 632 tonnes in 2004 ( FAO STAT, 2005 ). Live plant imports have increased over six-fold from 9550 to 62 800 tonnes in the same period ( AIPH, 1993; CBI, 2005 ). However, the

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alien native alien native alien native

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02468

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Figure 2 The numbers of plant pests establishing on non-native and native hosts, in relation to growth form of the host, for (a) all plant pests; expected pest numbers are derived from the proportional representation of aliens and natives in each growth form ( n = 131; missing data, n = 28; nonphytophages = 6); and (b) natural colonists; expected pest numbers are derived from the ratios of natives and non-natives in uncultivated habitats in the British flora ( n = 47, missing data, n = 1).


© 2007 The Authors


biggest proportionate increase in volume of imports has been seen with cut flowers, which have risen more than 150-fold, from 1330 to 201 764 tonnes during the 35-year period ( AIPH, 1993; HMCR, 2005 ). Phytosanitary regulations and their enforcement by the Plant Health services in the U.K. may be the key factor in suppressing the establishment rate of new pests, despite the increase in trade volume and variety.

Some uncertainty arises in the analysis of establishment rates because they are influenced by variations in recording (i.e. sampling) effort over time. Controlling for biases in ap-parent establishment rates is difficult because they may be due to changes in such factors as the number, ability and en-thusiasm of recorders, the availability of good identification keys, the number of taxonomists, the degree of general inter-est in non-native species and the accessibility of data. Indeed, the rate of accrual of species to national checklists may de-pend on the effort of key individuals for particular taxa. Ideally, future analyses will take recording effort into ac-count (e.g. by quantifying the volume of publications in rele-vant fields, or the number of records submitted to recording schemes). Comparing recent changes with historic trends is also problematic because it is often very difficult to catego-rize the native status of species. Thus, long-established non-natives may not be distinguished from natives, particularly in poorly-known taxa. As knowledge accumulates for taxa in a region, the probability of correctly identifying establishments should increase.

Establishment status

Natural colonists to Great Britain formed approximately one-third of all establishments, and they differed in taxonomic composition compared with human-assisted introductions. However, it is difficult to place this finding in a European context: colonists are often overlooked in appraisals of new arrivals, probably because data are known to be incomplete ( Kenis, 2005 ). It is likely that the natural spread of plant pests native to Europe, particularly from the Mediterranean region northwards, is a significant but little studied aspect of faunal change on the continent.

The fact that all natural colonizers were first detected in the southern regions of England, with East Kent heading the list of vice-counties (28%), was as expected, given that virtu-ally all colonizers had originated in mainland Europe. This pattern of colonization has been documented previously for migratory ( Bretherton, 1983 ) and colonizing ( Agassiz, 1996 ) British Lepidoptera, which formed the majority of such es-tablishments. It must be noted, however, that entomological recording effort is probably higher in the south of England, and here the climate is more suitable for species on the edges of their geographic ranges ( Asher et al. , 2001 ). Thus, prox-imity to source populations may not be the only determinant involved.

Recording effort is perhaps an important factor explaining why most human-assisted introductions were first detected in the south too (66%, mainly Surrey). Of these, 16 establish-ments were first detected at two botanical collections in

Surrey: the Royal Botanic Gardens at Kew, and the Royal Horticultural Society garden at Wisley. This is likely to be a consequence of the location of expertise in detecting pests that have spread to these sites, as well as these gardens act-ing as points of entry.

Introductions on ornamental plants

Ornamental plants accounted for 89% of all introductions and a similar high proportion has been found in France (Martinez & Malausa, 1999; Streito & Martinez, 2005 ), Italy ( Pelizzari et al. , 2005 ), and Serbia and Montenegro ( Glavendeki ć et al. , 2005 ). Ornamental plants currently pose a particular risk because they tend to be transported swiftly, with hitch-hikers intact, into suitable receptor habitats, and the volume of trade is increasing overall ( CBI, 2005 ). Coupled with this, it is possible that changing fashions in garden plants drive a regular turnover in plant species, and in the markets that supply them. In the U.K., for example, 60 – 70 000 types of plant (including cultivars and varieties) are available each year via nurseries, of which 4 – 5000 may be new to the market ( Smith et al. , 2006 ).

So far, all recent studies of introductions in Europe have identified Asia and North America as the major source of pests. Although no significant changes were detected in the sources of pests for the British data, in the last 10 years, 11 Asian species entered and established, suggesting that inva-sions from this continent may be about to increase. In Florida, establishments from Asia have increased subsequent to 1986 ( Thomas, 2000 ). However, careful examination of pathways is required to identify the relationships between trade and unintentional introductions. For example, although global trade has increased, the species richness or abundance of im-ported invertebrates may not be related in a straightforward way to the number of pathways or volume of trade; rather, particular commodities may account for a large proportion of arrivals in a pathway ( Work et al. , 2005 ).

Potential impacts of plant pests

The analyses of host plant use, and of the environments where pests established, showed that the potential impacts of pests were not exclusively on cultivated or uncultivated plants, except for those under cover. This considerable over-lap arose partly because many native hosts, or their close alien relatives, occur in both ornamental plantings (gardens, parks, etc.) and uncultivated habitats. For example, in domes-tic gardens, 50% of alien species nevertheless belong to na-tive genera ( Smith et al. , 2006 ), and relatively few herbivores are restricted to a single plant species. In addition, many pests were sufficiently broad in their host choice so as to be able to exploit cultivated and uncultivated plants (e.g. species of Acer , Pinus and Juniperus ).

Established pests, viewed overall, occurred disproportion-ately on alien compared with native woody hosts. This was most likely a consequence of human-assisted introductions being associated with ornamental plants, which include a higher than expected number of alien woody species ( Smith


© 2007 The Authors


et al. , 2006 ). Strikingly, however, when the hosts of natural colonists were compared with the ratios of naturalized alien and native plants in the British flora, there were many more colonists on woody aliens compared with perennials. It is un-clear whether woody species have more niches available for exploitation by colonists, or if the bias is an artefact of dif-ferential recording effort towards different growth forms.

More than 80% of natural colonists established in unculti-vated habitats. Colonists comprised mainly Lepidoptera, of which most were confined to a single host genus. Thus, the risks of natural colonists appear lower than human-assisted introductions; such species were dominated by the Homoptera (particularly scale insects and mealybugs), which had the highest proportion of oligophagous and polyphagous species.

Next steps

The assessment of impacts by non-native plant pests requires data on actual host damage and pest distribution. Both types of data will require considerable resources, for only a selec-tion of invaders have been systematically monitored for ex-ample Phyllonorycter leucographella and Phyllonorycter platani ( Nash et al. , 1994 ), Harmonia axyridis ( The Harlequin Ladybird Survey, 2006 ), Chrysolina americana and Lilioceris lilii ( RHS, 2006 ), although the Lepidoptera are already relatively well mapped.

In addition to filling in gaps for certain elements of the fauna, it is also important to develop the knowledge base in relation to new discoveries. Although the plant health serv-ices in the U.K. must be notified of new plant pests ( Cannon et al. , 2001 ), until the recent plans for national surveillance ( NNSS, 2007 ) and integrated monitoring across Europe ( DAISIE, 2007 ), there had been no standard mechanism for reporting information on all such species, or for establishing their status on an official list (e.g. compared with the British Ornithologist’s Union’s Records Committee; BOU, 2006 ). Reports of new discoveries are currently published in a range of expert, amateur journals, or in the grey literature, such as individual biological recording scheme newsletters. Historically at least, species have also been included in checklists without qualification as to whether they are estab-lished or just imported ( Malumphy, 2002 ). Therefore, a coor-dinated scheme for recording and assessing new plant pests would be desirable.

Plant health services in the U.K. have been relatively suc-cessful in preventing the establishment of quarantine inverte-brate pests during the last 35 years, through intercepting pests at points of entry and by eradicating outbreaks ( NAO, 2003 ). However, phytosanitary surveillance detected very few of the establishments reported in this paper, emphasizing the importance of expert amateurs and the public in assisting with surveillance. For example, many Leptinotarsa decemlin-eata are reported by the public each year ( Defra, 2006 ).

Conclusions

The present study provides a foundation for assessing recent plant pest establishments in Great Britain, and wider patterns

across continental Europe. However, a comprehensive assess-ment will only be possible once knowledge of certain taxa in the British fauna is more complete. Also, studies across Europe need to be better standardized before robust compari-sons can be made.

It may be true that once a non-native species has estab-lished, it is a priority to manage any subsequent economic and environmental damage ( Hill et al. , 2005 ). However, na-tional strategies to manage non-native organisms require both short- and long-term approaches. Arguably, the most impor-tant element of any national strategy must be to prevent es-tablishment, a lesson amply demonstrated by a wealth of invasive species research ( OTA, 1993; Defra, 2003 ). Thus, the analysis of plant pest establishments can identify gaps in phytosanitary risk assessment, and ultimately help suppress the introduction of non-native species.

Acknowledgements

We are grateful to Alan MacLeod (CSL) for information on trade statistics, to Lynne Matthews (ex-CSL) and Ellie Agallou (ex-CSL) for their assistance, and to Stéphane Pietravalle (CSL) for statistical advice. Mark Parsons (Butterfly Conservation) kindly responded to queries. Funding for this work was provided by the Plant Health Division of Defra.

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Accepted 10 April 2007


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Appendix

Summary plant pest data

Table A1 Summary plant pest data used for analyses of establishments in Great Britain, 1970 – 2004. Species are arranged alphabetically within classes

No. Name Year Pathway Status In-out Origin Principal hosts Sources

Nematoda

1 Paratrichodorus renifer Siddiqi

1990 Plants I Out Unconfirmed Rhododendron spp. 140

2 Pratylenchus bolivianus Corbett

1989 Plants I In S. America Alstromeria spp. 141, 142

3 Trichodorus sparsus Szczygiel

1973 – Plants I Out Unconfirmed Unconfirmed 141

4 Tylenchorhynchus claytoni Steiner

1990 Plants I Out Unconfirmed Rhododendron spp. 140

Platyhelminthes

5 Austaloplana ( Geoplana/ Caenoplana ) sanguinea Moseley var alba sensu Jones (1981)

1980 Plants I Out Australasia (An invertebrate predator)

144, a

6 Caenoplana purpurea Dendy

1986 Plants I In Australasia (An invertebrate predator)

145, b

Acari (Arachnida)

7 Aceria ficus Cotte 2000 Plants I In Cosmopolitan Ficus spp., Cudrania tricuspidata

1

8 Panonychus citri McGregor

1978 Plants I In Asia Citrus spp., Rosaceae, Moraceae

2, a

9 Pentarismus oregonensis

1978 Plants I Out N. America Cupressaceae a

10 Rhyncaphytoptus ficifoliae Keifer

2000 Plants I In Cosmopolitan Ficus spp. 1

11 Schizotetranychus celarius Banks

1995 Plants I Out Asia Sasaella masmunaena, Phyllostachys spp.

3, 4

12 Trisetacus chamaecypari Smith

2002 Plants I Out N. America × Cupressocyparis leylandii

5

13 Varroa destructor ( jacobsoni ) Oudemans

1992 Apiculture I Out Asia Apis mellifera (honey bee)

6, a

14 Vasates quadripedes Shimer

2002 Plants I Out N. America Acer spp 7, 8, 9

Coleoptera (Insecta)

15 Brachyderes incanus Linnaeus

1994 Plants I Out Europe Pinus nigra, Pinus spp. 10

16 Chrysolina americana Linnaeus

1994 Plants? I Out Europe Lamiaceae 11, 12, 13

17 Crypturgus subcribrosus Eggers

1986 Timber I Out Europe Picea abies 14

18 Dendroctonus micans Kugelmann

1973 Timber? I? Out Asia Picea abies, P. sitchensis

15, 16, 17

19 Diabrotica virgifera virgifera Le Conte

2003 Stowaway? I Out N. America Zea mays, Triticum 18

20 Diaprepes abbreviatus Linnaeus

1996 – Plants I In Americas Palmae 17, a

21 Harmonia axyridis Pallas 2004 Biocontrol Ii Out Asia (An invertebrate predator)

19

22 Henosepilachna argus Geoffroy

1997 Natural? C? Out Unconfirmed Bryonia dioica 20

23 Lixus scabricollis Boheman

1992 Natural? I i Out Europe Chenopodiaceae 21, 22

24 Luperomorpha xanthodera Fairmaire

2003 Plants? I Out Unconfirmed Rosa spp., Astilbe spp., Erysimum spp.

23, a

25 Otiorhynchus armadillo Rossi

1998 Plants I Out Europe Polyphagous 24, 25

Continued


© 2007 The Authors



26 Otiorhynchus aurifer Boheman

1978 Plants? I Out Unconfirmed Unconfirmed 25, 26

27 Otiorhynchus crataegi Germar

1985 Plants? I Out Europe Ligustrum 25

28 Otiorhynchus salicicola Heyden

2000 Plants I Out Europe Polyphagous 25, 27

29 Otiorhynchus setosulus Stierlin

1992 Plants? I Out Europe Unconfirmed 25, 28

30 Pachyrhinus mustela Herbst

1994 Plants I Out Europe Pinus nigra, Pinus spp. 10

31 Panspoeus guttatus Sharp

1981 ? I Out Australasia Unconfirmed 29

32 Scolytus laevis Chapuis 1981 Timber? I? Out Europe Ulmus spp. 30, 31 33 Scolytus pygmaeus

Fabricius 2000 Natural? C? Out Europe Ulmus spp. 32

Dictyoptera (Insecta)

34 Loboptera decipiens Germar

1997 Stowaway? I Out Europe Unconfirmed 33

Diptera (Insecta)

35 Braula schmitzi Örösi-pál 1994 Apiculture I Out Asia Apis mellifera (honey bee)

34

36 Contarinia quinquenotata Löw

1985 Plants I Out Europe Hemerocallis fulva 35

37 Dasineura gleditchiae Osten-Sacken

1983 Plants I Out N. America Gleditsia 36, 37

38 Phytomyza gymnostoma Loew

2002 ? I Out Europe Allium spp. 38

39 Phytomyza hellebori Kaltenbach

1997 Plants? I? Out Europe Helleborus spp. 39, a

40 Tephritis matricariae Löw 2000 Natural? C? Out Europe Crepis spp. 40 41 Tephritis praecox Löw 2002 Natural? C? Out Europe Calendula officinalis ? 41 42 Terellia fuscicornis Loew 2000 Plants? I? Out Europe Cynara scolymus,

Silybum marianum 42

Heteroptera (Insecta)

43 Brachycarenus tigrinus Schilling

2003 Natural C Out Europe Lepidium latifolium 43

44 Campylomma annulicornis Sig.

1978 Natural C Out Unconfirmed Salix viminalis 44, 45

45 Deraeocoris flavilinea A. Costa

1996 Natural C Out Europe Acer pseudoplatanus, Sambucus nigra

46

46 Eurydema ornata Linnaeus

1997 Natural C Out Europe Brassicaceae 47

47 Hypseloecus visci Puton 2003 Natural C Out Europe Viscum album L. 48 48 Liorhyssus hyalinus

Fabricius 1996 Natural C Out Europe Oleracea & Lactuca

spp. (Asteraceae) 44, 49, 50

49 Nezara viridula Linnaeus 2003 Natural? C? Out Cosmopolitan Polyphagous 51 50 Nysius senecionis

Schilling 1992 Natural C Out Europe Tripleurospermum/

Matricaria spp. 44, 52

51 Orsillus depressus Dallas 1987 Natural? C? Out Europe Cupressaceae 44, 53 52 Peritrechus gracilicornis

Puton 1977 Natural? C? Out Unconfirmed Unconfirmed 44, 54

53 Placochilus seladonicus Fall.

1977 Natural? C? Out Unconfirmed Knautia arvensis 44, 45

54 Stictopleurus abutilon Rossi

1996 Natural? C? Out Europe Asteraceae 44, 49, 50

55 Stictopleurus punctatonervosus Goeze

1998 Natural? C? Out Europe Asteraceae, mainly seeds

44, 49, 55

56 Tupiocoris ( Dicyphus ) rhododendri Dolling

1971 Plants? I Out N. America Rhododendron spp. 44, 56

57 Tuponia carayoni Wagner

1979 Natural? C? Out Europe Tamarix gallica 44, 57

Table A1 Continued

Continued


© 2007 The Authors



Homoptera (Insecta)

58 Acizzia uncatoides Ferris & Klyver

1990 Plants I In Australasia Acacia rivalis , A. retinoides

58

59 Acutaspis umbonifera Newstead

1970 – Plants I In Unconfirmed Unconfirmed a

60 Aleurothrixus floccosus Maskell

1986 Plants I In Americas Citrus spp. 59, a

61 Aonidia lauri Bouché 1990 Plants I Out Europe Laurus nobilis , Lauraceae

60

62 Aphis oenotherae Oestlund

1992 Plants? Ii Out N. America Oenothera spp. 61, 62

63 Appendiseta robiniae Gillette

1982 Plants? I Out N. America Robinia pseudacacia 62, 63

64 Bemisia afer Priesner & Hosny

1980 Plants I In – out Africa Laurus nobilis 64, 65

65 Cacopsylla fulguralis Kuwayama

2002 Plants I Out Asia Elaeagnus × ebbingei , E. commutata

66

66 Cinara (Cedrobium) lapportei Remaudière

1974 Natural C Out Africa Cedrus spp 67, 68

67 Cinara cedri Mimeur 1971 Plants? I? Out Europe Pinus nigra , Cedrus spp. 67 68 Cinara curvipes Patch 1999 Plants? I? Out N. America Cedrus atlantica 62 69 Crypturaphis grassii

Silvestri 1997 Plants? I? Out Europe Alnus cordata 62, 69

70 Eulecanium excrescens Ferris

1998 Plants I Out Asia Polyphagous 70, 71

71 Eulepidosaphes pyriformis Maskell

1975 Plants I Out Australasia Polyphagous 72

72 Eupulvinaria hydrangea Steinweden

1987 Plants I Out Asia Polyphagous 36

73 Fiorinia externa Ferris 1980 Plants I In Asia Conifers 36, 73 74 Geococcus coffeae

Green 1996 Plants I In Asia Polyphagous,

numerous families 74, a

75 Iassus scutellaris Fieber 1978 Natural? C? Out Unconfirmed Ulmus spp. 44, 75 76 Icerya purchasi Maskell 1996 Plants? I Out Australasia Polyphagous 76 77 Idiocerus ustulatus

Mulsant & Rey 1991 Natural? C? Out Europe Populus alba , P . ×

canescens 44, 77

78 Illinoia ( Masonaphis ) lambersi MacGillivray

1971 Plants I Out N. America Rhododendron sp. 62, 78

79 Illinoia ( Masonaphis ) liriodendri Monell

2003 Plants I Out N. America Liriodendron tulipifera , Magnolia spp.

a

80 Leucaspis podocarpi Green

1975 Plants I Out Australasia Podocarpus spp. 72

81 Livilla (Floria) variegata Löw

1978 Plants? I Out Europe Laburnum spp . 79

82 Macrosiphum albifrons Essig

1981 Stowaway? I Out N. America Lupinus spp . 36, 62, 80

83 Moritziella corticalis Kaltenbach

1970 Plants? I Out Unknown Quercus spp. 68, 69, 81

84 Myzus hemerocallis Takahashi

2000 Plants? I? Out Asia Hemerocallis spp., Agapanthus spp.

a

85 Myzus varians Davidson 1970 Plants? Ii Out Asia Clematis spp., Prunus persica

67, 82

86 Odonaspis greenii Cockerell

1970 – Plants? I In Asia Bamboos (Poaceae) 83, a

87 Paracolopha morrisoni Baker

1990 Plants? I Out Asia Bamboos a

88 Pentaloma nigra 1996 – Plants? I In Unconfirmed Unconfirmed a 89 Periphyllus

californiensis Shinji ? Plants? I? Out Asia Acer spp., occ.

Aesculus spp. 69, a

90 Phenacoccus defectus Ferris

1990 Plants I In N. America Echeveria spp., Sedum palmeri

74, 84

91 Pineus cembrae Cholodkovsky

1981 Plants? I? Out Europe Pinus cembra 68

92 Pineus similis Gillette 1971 Plants? I? Out N. America Picea sitchensis 68, 85

Continued

Table A1 Continued


© 2007 The Authors



93 Pinnaspis buxi Bouche ? Plants? I In Cosmopolitan Polyphagous 86, a 94 Planococcus

vovae Nasonov 1982 Plants? I Out Europe Cupressaceae 36, 87

95 Prociphilus fraxini Fabricius

1970 Plants? I? Out Europe Fraxinus excelsior , Abies alba

68, 69

96 Pseudaulacaspis dubia Maskell

1994 Plants? I Out Australasia Pteropsida (ferns) a

97 Rhizoecus aloes Williams and Pellizzari (1997)

1996 Plants? I In Unconfirmed Unconfirmed a

98 Russellaspis ( Asterolecanium ) pustulans Cockerell

1982 Plants I In Cosmopolitan Polyphagous 86, 88

99 Sitobian ( Macrosiphum) ptericolens Patch

1972 Plants I Out N. America Pteridium aquilinum 62, 89

100 Spilococcus cactearum Bouché

1988 Plants? I In Americas Cactaceae, Epadicaceae, Aizoaceae

74, 88

101 Stephanitis takeyai Drake & Mao

1995 Plants I Out Asia Pieris spp., Agarista populifolia

90

102 Takecallis arundicolens Clarke

? Plants I In – out Asia Bamboos a

103 Takecallis arundinariae Essig

1999 Plants I In – out Asia Phyllostachys aurea 91, a

104 Takecallis taiwanus Takahashi

1999 Plants I In – out Asia Bamboos 91

105 Tinocallis nevskyi Remaudière, Quednau & Heie

1995 Plants? I? Out Asia Ulmaceae, Ulmus spp. 62, 92

106 Trichosiphonaphis polygonifoliae Shinji

1999 Plants? I? Out Asia Lonicera tartarica , Polygonum spp.

62

107 Trioza vitreoradiata Maskell

1993 Plants I Out Australasia Pittosporum tenuifolium , P. crassifolium

93

108 Trochiscococcus speciosus De Lotto

1996 Plants? I In Unconfirmed Unconfirmed a

109 Uroleucon erigeronensis Thomas

1973 Plants? I? Out N. America Conyza canadensis , Asteraceae

62

110 Utamphorophora humboldti Essig

1974 Plants? I? Out N. America Physocarpus opulifolius , Poaceae

62, 67

111 Vryburgia (Chorizococcus) brevicruris McKenzie

1978 Plants? I In N. America Asclepiadaceae & Cactaceae

87

112 Vryburgia amaryllidis Bouché ( Chorizocccus lounsburyi)

1970 – Plants? I In – out Unconfirmed Liliaceae 94, a

113 Wahlgreniella nervata Gillette

1973 Plants? I? Out N. America Rosa spp. & cultivars , Arbutus spp.

62, 67

Hymenoptera (Insecta)

114 Andricus corruptrix Schlechtendal

1972 Plants? I? Out Europe Quercus cerris 95, 96

115 Andricus grossulariiae Giraud

2000 Plants? I? Out Europe Quercus cerris 96

116 Andricus lignicola Hartig 1972 Plants? I? Out Europe Quercus cerris , Q.robur

95, 96, 97

117 Aphelonyx cerricola Giraud

1993 Natural? C? Out Europe Quercus cerris

96, 98

118 Arge berberidis Schrank 2000 Plants I Out Europe Berberis spp., Mahonia spp.

99

Lepidoptera (Insecta)

119 Argyresthia cupressella Walsingham

1997 Plants I Out N. America Juniperus spp., Cupressaceae

100, 101

120 Argyresthia trifasciata Staudinger

1982 Plants I Out Europe Juniperus spp., Cupressaceae

102, 103

121 Athrips rancidella Herrich-Schäffer

1971 Plants? I? Out Europe Cotoneaster horizontalis 101, 104, 105

Continued

Table A1 Continued


© 2007 The Authors



122 Caloptilia rufipennella Hübner

1970 Natural C Out Europe Acer pseudoplatanus 103, 106, 107

123 Cameraria ohridella Deschka & Dimi ć

2002 Stowaway? I? Out Europe Aesculus hippocastanum

108, a

124 Clostera anachoreta Denis & Schiffermüller

1979 Natural C Out Europe Salix spp. & Populus spp.

101, 109

125 Cochylis molliculana Zeller 1993 Natural C Out Europe Picris echioides 101, 105, 110 126 Coleophora aestuariella

Bradley 1981 Natural C Out Europe Suaeda maritima 101, 111, 112

127 Coleophora fuscicornis Zeller

1973 Natural C Out Europe Vicia tetrapserma 101, 112

128 Coleophora linosyridella Fuchs

1978 Natural C Out Europe Aster tripolium 101, 112

129 Cydia illutana Herrich-Schäffer

1975 Plants? I? Out Europe Picea abies , Abies alba , Larix decidua

101, 113

130 Cydia medicaginis Kuznetsov

1970 Natural? C? Out Europe Medicago spp. 101, 105, 114

131 Dioryctria schuetzeella Fuchs

1980 Natural C Out Europe Picea abies 101, 105

132 Dioryctria sylvestrella Ratzeburg

1999 Natural C? Out Europe Pinus spp. , Picea spp. 101, 115, 116

133 Dryobota labecula Esp. 1999 Natural C Out Europe Quercus ilex 117, 118 134 Duponchelia

fovealis Zeller 1996 Plants I In – out Europe polyphagous on

cultivated plants 105, 119

135 Ectoedemia (Etainia) sericopeza Zeller

1975 Natural? I? Out Europe Acer platanoides , A.campestre ?

101

136 Ectoedemia erythrogenella Joannis

1973 Natural C Out Europe Rubus fruticosus 101, 120

137 Ectoedemia hannoverella Glitz

2003 Plants? I? Out Europe Populus × canadensis 121

138 Ectoedemia heringella Mariani

2002 Plants? I? Out Europe Quercus ilex 122, 118

139 Emmetia heinemanni Wocke

1984 Natural? C? Out Europe Rubus spp. 101, 118, 148

140 Eucosma metzneriana Treitschke

1998 Natural C? Out Europe Artemeria absinthium , A. vilgare

101, 105

141 Eupithecia ultimaria Boisduval

1989 Natural C Out Europe Tamarix gallica 101, 109

142 Evergestis limbata Linnaeus

1993 Natural C Out Europe Alliaria petiolata, Sisymbrium officinale

101, 105, 123

143 Gelechia sabinellus Zeller 1971 Plants I Out Europe Juniperus spp. 105, 124, 125 144 Gelechia senticetella

Staudinger 1988 Plants I Out Europe Juniperus spp.,

Cupressaceae 101, 125, 126

145 Haimbachia cicatricella Hübner

1999 Natural C Out Europe Schoenoplectus lacustris

101, 105

146 Hecatera dysodea Denis & Schiffermüller

1997 Natural C Out Europe Lactuca spp. 101, 105

147 Hypena obsitlis Hübner 1990 Natural C Out Europe Parietaria judaica 101, 105, 127 148 Lampronia flavimitrella

Hübner 1974 Natural? C? Out Europe Rubus spp.,

inc. R. idaeus 101, 128

149 Monochroa moyses Uffen 1971 Natural C Out Europe Bolboschoenus maritimus

101, 125, 129

150 Monochroa niphognatha Gozmány

1984 Natural C Out Europe Persicaria amphibia? 101, 104, 125

151 Peribatodes secundaria Esper

1981 Natural C Out Europe Picea abies , Pinaceae , Cupressaceae

101, 105

152 Phyllocnistis xenia Hering 1974 Natural C Out Europe Populus × canescens 101 153 Phyllonorycter

leucographella Zeller 1989 Plants? I? Out Europe Pyracantha , Malus ,

Sorbus , Crataegus 103, 105, 130

154 Phyllonorycter platani Staudinger

1990 Plants? I? Out Europe Platanus × hispanica 101, 105, 131

155 Sciota adelphella Fischer von Röslerstamm

1992 Natural C Out Europe Salix alba 101, 105

Continued

Table A1 Continued


© 2007 The Authors



156 Scythris inspersella Hübner

1977 Natural? C? Out Europe Chamerion angustifolium

105, 132, 133

157 Thera cupressata Geyer 1984 Natural C Out Europe Cupressus spp . 101, 109, 134 158 Tinagma balteolella

Fischer von Röslerstamm

1975 Natural C Out Europe Echium vulgare 101, 105, 135

159 Vitula biviella Zeller 1997 Natural C Out Europe Pinus spp. 118, 136

Phasmatodea (Insecta)

160 Bacillus rossius Rossi 1986 Release I Out Unconfirmed Unconfirmed 33, 143 161 Sipyloidea sipylus

Westwood 1986 Release I Out Unconfirmed Unconfirmed 33, 143

Thysanoptera (Insecta)

162 Echinothrips americanus Morgan

1995 Plants I In N. America Polyphagous 146

163 Frankliniella occidentalis Pergande

1986 Plants I In N. America Polyphagous 147

Mollusca

164 Boettgerilla pallens Simroth

1972 Plants? I Out Europe Polyphagous 137, 138, 139

Year, year of first recorded establishment ; – , indicates that establishment was sometime after that date. Pathway, means of entry, where ‘plants’ refers to trade, and ‘natural’ indicates natural colonists. Status: ‘I’, human-assisted introductions; ‘i’, indirectly introduced (i.e. natural arrivals that had been initially introduced elsewhere); ‘C’, natural colonists. In – out, species established indoors, outside, or both. Origin: ‘Neotropics’, refers to species of tropical America or the Caribbean. ‘?’, denotes an element of uncertainty; ‘a’, refers to unpublished CSL files.

Literature sources to plant pest data used in analyses 1. Ostojá-Starzewski, J.C. (2002) Aceria ficus (Cotte) and Rhyn-

caphytoptus ficifoliae Keifer (Acari: Eriophyoidea) first records in the British Isles, British Journal of Entomology and Natural History , 15 , 79 – 82.

2. Cheek, S. & Baker, R.H.A. (1993) Panonychus citri , the Citrus Red Mite. Plant Pest Notice No. 16. Central Science Labora-tory (MAFF), U.K.

3. Ostojá-Starzewski, J.C. (2000) Schizotetranychus celarius (Banks, 1917) (Acari: Prostigmata). A mite pest of bamboo; first records for Britain and two new host records, British Jour-nal of Entomology and Natural History , 13 , 95 – 97.

4. Stapleton, C. (1996) Mite alert. The Bamboo Society Newslet-ter, p. 24.

5. Ostojá-Starzewski, J.C. & Halstead, A.J. (2006) Trisetacus chamaecypari Smith (Acari: Eriophyoidea: Phytoptidae) new to Britain, damaging × Cupressocyparis leylandii (Dallimore & Jackson) (Cupressaceae) a new host for this mite, British Journal of Entomology and Natural History , 19 , 201 – 205.

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22. Morris, M.G. (2002) True weevils (part 1), Coleoptera: Curculion-idae (Subfamilies Raymondionyminae to Smirconychinae). Hand-books for the Identification of British Insects, Vol. 5 (17b). Royal Entomological Society, U.K. and Field Studies Council, U.K.

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27. Cole, S. (2002) 2001 Annual Exhibition, Imperial College, London SW7 – November 2001. Coleoptera, British Journal of Entomology and Natural History , 11 , 176.

28. Morris, M.G. (1997) Broad-nosed weevils Coleoptera: Curcu-lionidae (Entiminae). Handbooks for the Identification of Brit-ish Insects, Vol. 5 (17a). Royal Entomological Society, U.K.

29. Owen, J.A., Allan, A.A., Carter, I.S. & von Hayek, C.M.F. (1985) Panspoeus guttatus Sharp (Col., Elateridae) new to Britain, Entomologist’s Monthly Magazine , 121 , 91 – 95.

30. Atkins, P.M., O’Callaghan, D.P. & Kirby, S.G. (1981) Scolytus laevis (Chapuis) (Coleoptera, Scolytidae) new to Britain, Ento-mologist’s Gazette , 32 , 280.

31. Winter, T.G. (1991) Interceptions of exotic bark beetles (Col., Scolytidae) on timber imports into Great Britain, 1980 – 1988, Entomologist’s Monthly Magazine , 127 , 13 – 17.

32. Heal, N.F. (2003) Scolytus pygmaeus (Fabricius, 1787) (Scolyt-idae) – a new arrival to Britain, The Coleopterist , 12 , 57 – 60.

33. Marshall, J.A. (2001) Grasshoppers, crickets & allies. The Changing Wildlife of Great Britain and Ireland, Systematics Association Special Volumes Series 62 (ed. by D. L. Hawksworth), pp. 328 – 339. Taylor and Francis, U.K.

34. Dobson, J.R. (1998) A ‘bee louse’ Braula schmitzi Örösi-pál (Diptera: Braulidae) new to the British Isles, and the status of Braula spp. in England and Wales, British Journal of Entomol-ogy and Natural History , 11 , 139 – 148.

35. Halstead, A.J. & Harris, K.M. (1990) First British record of a gall midge pest of day lily ( Hemerocallis fulva ), British Jour-nal of Entomology and Natural History , 3 , 1 – 2.

36. Halstead, A.J. (1992) The 1991 Presidential Address – Part 2. Some horticultural pests new to Britain in recent years, British Journal of Entomology and Natural History , 5 , 41 – 47.

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39. Spencer, K.A. (1976) The Agromyzidae (Diptera) of Fen-noscandia and Denmark. Fauna Entomologica Scandinavica, Vol. 5. Scandinavian Science Press, Denmark.

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42. Whittington, A.E. (2002) Terellia fuscicornis (Loew, 1844) (Dipt., Tephritidae) new to Britain, Entomologist’s Monthly Magazine , 138 , 119 – 120.

43. Jones, R.A. (2004) Brachycarenus tigrinus (Schilling) (Hemip-tera: Rhopalidae) new to Britain, British Journal of Entomol-ogy and Natural History , 17 , 137 – 141.

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45. Nau, B.S. (1979) Two plant bugs new to Britain, Placochilus seladonicus (Fall.) and Campylomma annulicornis (Sig.) (Het-

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46. Miller, D.J.P. (2001) Deraeocoris flavilinea (A.Costa) (Hemip-tera: Miridae), new to Britain, British Journal of Entomology and Natural History , 14 , 133 – 136.

47. Nau, B.S. (2006) Eurydema ornatum (L.) living in the south of England. Heteroptera Newsletter, no. 6, Autumn 2005, p. 15.

48. Gibbs, D. & Nau, B. (2005) Hypseloecus visci (Puton) (Hemip-tera: Miridae) a mistletoe bug new to Britain, British Journal of Entomology and Natural History , 18 , 159 – 162.

49. Southwood, T.R.E. & Leston, D. (1959) Land and Water Bugs of the British Isles. Warne, U.K.

50. Nau, B.S. (1997) Range-changes in some Hemiptera-Homop-tera in Bedfordshire, Entomologist’s Monthly Magazine , 133 , 261 – 262.

51. Shardlow, M.E.A. & Taylor, R, (2004) Is the Southern Green Shield Bug, Nezara viridula (L.) (Hemiptera: Pentatomidae) another species colonising Britain due to climate change? Brit-ish Journal of Entomology and Natural History , 17 , 143 – 146.

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