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Zeitschrift/Journal: Bonn zoological Bulletin - früher Bonner Zoologische Beiträge.

Jahr/Year: 2005/2006

Band/Volume: 54

Autor(en)/Author(s): Beenen Ron

Artikel/Article: Translocation in Leaf Beetles (Coleoptera: Chrysomelidae) 179-199 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at

Bonner zoologische Beiträge Band 54 (2005) Heft 4 Seiten 179-199 Bonn, Oktober 2006

1 Translocation in Leaf Beetles (Coleóptera: Chrysomelidae)

i Ron Beenen Nieuwegein, The Netherlands

Abstract. An overview of leaf beetle translocations, and the possible effects related to endangerment of native leaf bee-

tle species is presented. Translocation is defined as the movement of living organisms from one area to another across natural barriers. In the new area the translocated organism lives free. Translocations can be intentional or accidental.

The database of records of translocated leaf beetles is compiled using published records as the main source of information. In this study 126 leaf beetle species have been recorded to have been translocated at least once. Most translocated species occur in the subfamilies Chrysomelinae, Galerucinae, Alticinae, Cassidinae and Hispinae. Most unintentional successful translocations occur in mono- and oligophagous species that develop on cultivated and non-indigenous plant species. At present, no negative effects of translocated Chrysomelidae on biodiversity have been reported. The successful control of imported weeds by introduced Chrysomelidae can be seen as a positive effect to biodiversity since some of these weeds prevent natural processes in ecosystems. Historical distribution patterns of Chrysomelidae are discussed in relation to the translocations.

Keywords. Coleóptera, Chrysomelidae, invasive species, bioinvasion, alien species, translocation, introduction

1. INTRODUCTION phagous species (including Chrysomelidae) exists (e.g., Gonzáles-Megías & Gómez (2003)). One of the major threats to native biological diversity is now acknowledged by scientists and governments to be When a population of a non-indigenous species out biological invasions caused by alien invasive species competes a population of a native species, the invasive (IUCN 2000, Union of Concerned Scientists 2001). species must have reached a population level in which

This is not a new item: the threat of invasive species for its competition can be effective. When the population the conservation of biodiversity was already pointed out level is too low, it is unlikely that there will be a large by ELTON in 1933 (cf. SlMBERLOFF's foreword in effect on the population of the indigenous species. Co-

ELTON 2000). Meanwhile it is supposed that every year lautti & MacIsaac (2004) developed a framework for a wide range of alien species are imported into many defining different stages during invasions. After the new countries through international trade both intentionally species is established a non-indigenous species may be as trade products (horticulture, pets, etc.) or uninten- localised and rare (a), widespread but rare (b) or wide- tionally (for example by hitch-hiking on legitimate spread and dominant (c). In this last mentioned stage, products). Little is known about the number of leaf bee- out competing of indigenous species may be the result. tles (Coleóptera, Chrysomelidae) that are translocated in Therefore, for our purpose, we need to know whether this way or about their impact on biological diversity. translocated Chrysomelidae have become established.

Translocated species can be a threat in the way that a That is the first step. Subsequently we would like to native leaf beetle is displaced by an alien leaf beetle know the status of the established species. The aim of species. On the other hand translocated species can re- this study is to get an overview of leaf beetle introduc- duce indigenous plant species and thus affect the native tions, and the possible effects related to endangerment herbivores depending on it. In this study we restrict con- of native leaf beetle species. sideration to leaf beetle translocations and their effects Natural range expansion of a species, as for example re- on native leaf beetle populations caused by competition. corded for Chaetocnema major Jacquelin-Duval by Interspecific competition is any interaction between two DÖBERL (1994a), is not treated here. Here we consider or more species populations which adversely affects all transport across natural barriers as translocation. their growth and survival (Odum 1971). The importance Both intentionally (e.g., in biological control programs) of interspecific competition is not clear (STEWART as unintentionally. The IUCN (1987) defines transloca- 1996) but evidence for competition among phyto- tion more strictly and restricts to "introductions", "reintroductions" and "re-stocking". Introductions can be effective tools in the management of natural and man

,h made environments. Re-introduction and re-stocking as Paper presented to the 6 International Symposium on the Chry-

somelidae, Bonn, Germany, May 7, 2004. an aim in nature management has not been used with .

Chrysomelidae species. Thus, for this purpose we do not (after several unsuccessful attempts from 1876). The have to include the IUCN-restrictions. beetle spread rapidly in Europe despite intensive opera-

tions to control it. It was first reported in Belgium and Here we define translocation as the movement of living Spain in 1935,' Luxembourg in 1936, the Netherlands organisms from one area to another across natural barri- and Switzerland in 1937, Austria in 1941, Hungary and ers. In the new area the translocated organism lives free. the former Czechoslovakia in 1945, Poland and Roma- Translocations can be intentional or accidental. The role nia in 1947, and Turkey in 1949 (CAB-CPC 2003). In of man is not always clear in cases of translocation. Appendix 1 the year 1922 is taken as the moment of es-

tablishment in Europe although it has been found earlier 2. METHODS in France and established many years later in some The database of records of intentionally or non- European countries. intentionally introduced leaf beetles is compiled using On the other hand a species may not have established in published records as the main source of information. all parts of the country to which it is translocated. For Because not all published information was available for example: all attempts to establish L. jacobaeae in On- this study there is a chance that information is lacking; tario, New Bainswick and Prince Edward Island were this is believed to be compensated by other publications unsuccessful. In Canada it is at present established only in which similar information is available. in British Columbia. In the United States it is currently known from California and Oregon Only the traditional leaf beetles have been studied, i.e. (LeSage 1988). exclusive the Bruchidae. Subfamily names used in this study are the traditional as have been used by Seeno & 3. RESULTS WILCOX (1982). All species group names of Chrysomeli- In this study 126 leaf beetle species have been recorded to dae in Appendix 1 are scientific names complete with au- have been translocated at least once. They are listed in thor. In the text they are without author names, except for Appendix 1. Because many species have been translo- those species that are not listed in Appendix 1 cated more than once, a total number of 230 translocation

A translocation event is defined as a translocation from events in Chrysomelidae is listed (Appendix 1). Of each an area where the species is indigenous to an area where event information is given on place of origin of the leaf the species previously did not occur. In this respect the beetle, place where it is translocated to, moment of trans- translocation of Chrysolina hyperici from England to location, degree of success, host plant and source (refer-

Australia in 1930 and from France to Australia in 1980 ence). In Figure 1 the length of each bar represents the to- are considered as two different translocation events. In tal number of species of each subfamily. Most the case of biocontrol, translocations with different tar- translocated species occur in the subfamilies Chrysomeli- gets (host plants) are considered as different transloca- nae, Galerucinae, Alticinae, Cassidinae and Hispinae. In tion events. Figure 2 the length of the bar represents the number of translocations of each subfamily. The bar for the subfam- Excluded from the database are natural expansions. ily Hispinae is much larger than in Figure 1, indicating However we have to admit that it is not always clear to that some species have been translocated several times. discriminate between natural range expansion and transport across natural barriers. This might be the case Translocations can be intentional or unintentional. The in species that are still in the phase of post-glacial ex- number of intentional translocations exceeds the number pansion. of unintentional translocations (Table 1 ). This is not surprising. Unintentional translocations are hard to de- Also accidental translocations that have resulted in es- tect; they mostly have not been published. Publications tablishment in artificial environments as in greenhouses on species that have been unintentionally translocated are not included in this research. In the Netherlands for mostly will deal with species that have become estab- example the alticine Acrocrypta purpurea Baly has es- lished in the newly inhabited land. When established tablished in a greenhouse where during several genera- they are more likely to be observed. Of the unintentional tions it damaged orchids (DÖBERL 1994b). Acroaypta translocated species 81 % (70 out of 86) have become purpurea originates from South East Asia and it is established against 63 % (91 out of 144) of the inten- unlikely that it will survive in the Netherlands outside tionally translocated species. The successes of inten- green )houses. ( tional translocations that are part of a biological control

Sometimes it is difficult to define the exact moment of program are monitored. This means that not only infor- establishment. Leptinotarsa decemlineata, for example, mation on successful translocations are available but was discovered in the Rocky Mountains and described also of the unsuccessful releases. On the other hand the in 1824. Suddenly, in 1859, it began devastating potato intentional translocations mostly take place only after crops 100 miles west of Omaha, Nebraska, USA. Over intensive study. In biological control programs the suc- the next few years, the beetle spread eastward to the At- cess (and the risk) of the introduction is carefully stud- lantic coast, which it reached in 1874. Leptinotarsa de- ied before release. In that respect it is surprising to no- cemlineata became established in Europe following its tice that 27 % (39 out of 144) of the intentionally introduction from the USA to Bordeaux, France in 1922 translocated species have not become established. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Ron Beenen: Translocation in Leaf Beetles [81

most leaf beetles, host plants from the same genus or family have to be available in the new country.

It is interesting to find out what types of plants these successfully unintended translocated species feed on. Four categories of plants species have been distin- guished: cultivated plant species, non-indigenous "wild" plants, indigenous "wild plants" and a last category of

which information was insufficient (Fig. 4). It is evident that the majority of unintended successful translocation of leaf beetles occurred on cultivated plants and on non- indigenous (introduced plant species). Only a minority occurs on plants that are indigenous. Fig. 1. Results of translocations in Chrysomelidae. Number of translocated species per subfamily. Subfamily abbreviations: CRIO = Criocerinae; ZEUG = Zeugophorinae; CRYP = Cryptocephalinae; EUMO = Eumolpinae; CHLA = Chlamisi- nae; = = Galerucinae; = CHRY Chrysomelinae; GALE ALTI species = = Alticinae; CASS Cassidinae; HISP Hispinae. Results: UE O events = unintentional translocation followed by establishment; UN = unintentional, not established; UD = unintentional, establishment doubtful; IE = intentional and established; IN = intentional and not established: ID = intentional and doubtful.

oligo poly

Fig. 3. Number of unintentionally translocated leaf beetles that have established per categories of food plants. Mono = monophagous, oligo = oligophagous, poly = polyphagous. Species = number of translocated species, events = number of translocation events.

Fig. 2. Results of translocations in Chrysomelidae. Events pet-

subfamily. Subfamily abbreviations as in Figure 1. Results: UE = unintentional translocation followed by establishment; UN = unintentional, not established; UD = unintentional, establishment doubtful; IE = intentional and established; IN = non-indig indigen intentional and not established; ID = intentional and doubtful.

Fig. 4. Number of unintentionally translocated leaf beetles that Intentionally translocated species mostly have very re- have established per plant category: cultivated plants, non- stricted food preference. The chances that they become indigenous plants, indigenous plants and plants of which this information is a problem as a result of feeding on non-target species is not available. lowest in monophagous species. We analysed the food preference of the unintentionally translocated species. In 4. MISCELLANEOUS RESULTS Figure 3 the number of species is presented for the next

groups of phytophages: monophages - species that de- Apart from the results described above, interesting in- velop on a single plant species or genus; oligophages - formation became available on a variety of subjects. An species that develop on more than one plant genus of a extract of this is presented in this section. The following single plant family; polyphages - species that develop subjects are presented: description of species based on on plant species of more than one family. It shows that translocated specimens; unused biological control most of these successful translocated species are mono- agents; the use of indigenous species in biological con- or oligophagous; they develop on plant species within a trol; anecdotal information on the ways unintentional single genus or within a single plant family. This more translocations have taken place and finally non-target or less implies that, because of the food restriction of effects. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 182 Bonner zoologische Beiträge 54 (2005)

4.1. Description of species based on translocated establish (JULIEN 1992). In order to control Calystegia specimens sepium, native organism of Chirida guttata have been collected in Ontario in 1971 and released in British Co- Some species description were based on displaced lumbia. They did not establish either (Julien 1992). The specimens: Falsoplatyxantha diversicornis Pic, Lupe- same holds true for Metriona purpurata native organism rus marginalis Allard and Aulacophora pannonica Csiki collected in 1979 in Saskatchewan and released in Al- (= Hoplasoma unicolor 111.) (all Galerucinae). The type berta in an attempt to extend its range and control Con- specimen(s) of Falsoplatyxantha diversicornis were volvulus arvensis (JULIEN 1992). However, Alfica fovei- from Turkey, but the author already stated that the spe- collis Jacoby (Alticinae) a native species of Thailand cies most probably originally came from the Indies was used in this country to control Ludwigia adscen- ("Acquis comme provenant d'Asie Mineure, mais origi- dens. It caused considerable damage with satisfactory naire vraisemblablement des Indes") (Pic 1931). Of degree of control (JULIEN 1992). Gastrophysa atro- Hoplasoma unicolor a single specimen was collected in cyanea a native species introduced in Noto Peninsula Hungary and described as Aulacophora pannonica by (Japan) to control Rumex obtusifolius quickly estab- CSIKI (1953). SlLFVERBERG (1978) studied the type lished, increased its population and spread rapidly. specimen of A. pannonica and reduced it to a junior Population of the beetle reached satisfactory levels synonym of H. unicolor. The type specimen of A. pan- within four years of release (Julien 1992). Metriona bi- nonica was probably imported from the Oriental region. color, a native organism collected in 1971 in Ontario Although there always remains the possibility of misla- and released in British Columbia in order to control Ca- belled specimens (see MOHAMEDSAID 2001). Luperus lystegia sepium, established. In 1985 it was found 14 marginalis was described by Allard (1890) from the km from the release site (JULIEN 1992). Falkland archipelago and according JOLIVET & VERMA

(2002) evidently imported there in historical times. Both Leptinotarsa defecto and L. texana are indigenous Apart from L. marginalis no Chrysomelidae have been in the USA (Texas) and have been proposed as a bio- mentioned from the Falkland archipelago. logical control agent against three invasive Solanum species. The results of the study suggest that of the in-

vasive Solanum species, 5. torvum may be included in 4.2. Unused biological control agents the potential host range of L. texana. It is suggested that

Some species have been selected to be used as a bio- this "new association" approach might result in a sub- logical control agent but have not been used yet: Agen- stantial control of one of Florida's most invasive so- iosa electoralis Vogel (Chrysomelinae) was selected to lanaceous weeds with acceptable ecological risks (CUDA be introduced in Australia to control Chrysanthemoides et al. 2002). monilifera (Jolivet 2001). Charidotes pygmaea Klug (Cassidinae) was selected to be released in Australia to Native biological control agents offer potential advan- tages over non-native agents, because they may have lit- control species of Lantana (Jolivet 2001 ). Homichloda barkeri Jacoby (Alticinae) from Kenya was selected to tle impact on non-target native species that have coex- control Acacia nilotica in Queensland (Jolivet 2001). isted with the control agent (SHELDON et al. 1995). Trachyaphthona sórdida (Baly) (Alticinae) a specialist herbivore on Paederia foetida (Skunk Vine) in China 4.4. Registered unintentional translocations and Japan, which is a very invasive weed in Florida rarely aspects unintentional translocations spreading in other parts of the Southern Unites States. Very of have registered. In a few cases they can however be re- Trachyaphthona sórdida is of special interest to be be been used as a biological agent (Pemberton & Pratt 2002). constructed. Sometimes unintentional introductions have been the result of intentional introduction during a Several species have been selected for the control of in- biological control program. Aphthona nigriscutis was vasive plants but finally introduction has been aban- accidentally released in Canada in 1982 together with doned because the species proved to not selective be Aphthona cyparissiae. The release site was treated with in their choice. is the case for enough food This exam- herbicide and insecticide in order to eradicate this spe- ple for Platyphora biforis (Germar), P. conviva (Stál), P. cies (Julien 1992). And Chrysolina hyperici is believed (Stál) P. for the nigronotata and paraguana (Jacoby) to have been introduced accidentally in Hawaii together control of Solanaceae (Jolivet 2001). with C. quadrigemina in 1965. It was discovered in 1970 (Julien 1992) and Jolivet (2001) assumes this

4.3. The use of indigenous species in biological species is established in Hawaii. In 1979 a Longitarsus control species referred to as L. jacobaeae has been released in Australia from France. This species more closely re- Very rarely indigenous species are successful in control- sembles L. flavicornis. It is established and spreading, ling nuisance herbs. Both Chelymorpha cassidea (F.) causing high reduction in weed density at some sites (Cassidinae) and Chirida guttata (Olivier) (Cassidinae) (Julien 1992). have been collected in 1979 in Saskatchewan and released in Alberta in an attempt to extend their ranges for Longitarsus ganglbaueri, a European species, is re- the control of Convolvulus arvensis, where they did not cently found in Oregon. LeSage (1988) assumed it to © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Ron Beenen: Translocation in Leaf Beetles 1X3

be accidentally introduced with those of L. jacobaeae %]). The observed representation of subfamilies among imported from Italy and introduced in Oregon. the translocated species seems to be a mix of the subfamily representations in these biogeographical regions. Xanthogaleruca luteola was accidentally introduced For exampled, in the Palearctic Eumolpinae are repre- into Britain from Bolzano (Italy) in July 1986. Several sented by a low number of species. specimens had accidentally been transported in camping single equipment (SMITH 1990). The specimen recorded It is most likely that the number of unintentional trans-

by STERRENBURG (1989) for the Netherlands may also locations is highly underestimated. Unintentional trans-

have been based on a displaced specimen. This species locations that did not result in establishment are rarely has been recorded for the Netherlands only from the detected and rarely published. However, unintentional very south of the country (Beenen 1998) translocations that resulted in establishment and showed negative effects are likely to have been noticed. Such 4.5. Non-target effects translocations are of importance for our main question, namely the impact of translocations on the biodiversity. Zygogramma bicolorata is an efficient biocontrol agent that can have significant negative impact on the growth Sometimes the reason why an introduced species has

and reproduction of Parthenium hysterophorus (Dhile- not established is clear. Alfica carduorum was intro- EPAN et al. 2000). Zygogramma bicolorata breeds in In- duced in Canada in 1963 from Switzerland and France scale dia on small on Xanthium strumarium and feeds to control Cirsium arvense. It did not establish in Al- on Helianthus annuus. Feeding on H. annuus is re- berta, British Colombia, Nova Scotia or Ontario. Slow garded unwanted non-target feeding. Parthenium pol- development in cool summers exposed larvae to high len, which contains parthenin, when deposited on Heli- predation (JULIEN 1992). It was also introduced to Great anthus annuus, makes the latter attractive to beetles in Britain in 1969 and 1970 from France. Several thou- absence of Parthenium. Continuous feeding on Helian- sands of specimens were released at Silkwood Park, As- thus annuus retards ovarian development and thus af- cot, Berkshire and at three sites west of Cardiff, fects et al. fecundity (VlRAKTAMATH 2004). Glamorganshire, South Wales (Baker et al. 1972, Co.X

2000). They survived the winter in cages only. The cli- Julien (1992) mentioned the case of Chrysolina hy- mate is too cold and wet to allow survival, except lo- perici that was introduced in New Zealand in 1947 from cally (JULIEN 1992). Chrysolina varians was introduced England to control Hypericum perforatum but attacked in Australia from England in 1930 to control Hypericum Hypericum androsaemum. Despite some early damage perforatum and did not establish. It is considered to the insect has not persisted on H. androsaemum and has have suffered from heavy predation and unfavourable not established. In this case the non-target effect did not climate (JULIEN 1992). It was introduced in Canada persist. Galerucella calmariensis and Galerucella pu- from Sweden in 1957 to control Hypericum perforatum silla have been introduced to North America in 1992 as and did not establish in British Columbia. Release sites a biological control agent for Lythrum salicariae and are were too dry during summer (JULIEN 1992). Adults of now established in many US states and Canadian prov- the alticinae Disonycha glabrata (F.) have been released inces. At some sites short-term attack on Rosa multi- to control Amaranthus retroflexus from Massachusetts flora, Potentilla anserina and Decodon verticillatus has (U.S.A.) in Red River Valley, North Dakota in 1979 and been observed. This "spillover" does not constitute a 1980 but failed to overwinter (Julien 1992). Longitar- host shift since the beetles are unable to complete de- sus albineus was introduced in Australia in 1979 and velopment on these non-target plants (BLOSSEY et al. 1981 from Greece and France to control Heliotropium 2001). europaeum. Establishment failed due to drought, which eliminated the host (JULIEN 1992). Physonota alutacea 5. DISCUSSION was introduced in Mauritius from Trinidad in 1947 to

control Cordia curassavica. It did not establish due to It is difficult to give an explanation of the observed rela- interference by ants (Julien 1992). tive high number of species from the subfamilies Chry- somelinae, Galerucinae, Alticinae, Cassidinae and The observed low percentage of polyphagous species (7 Hispinae that have been translocated. It cannot be ex- %) that have established after unintentional transloca- plained from the representation of these subfamilies tions is not surprising. It seems to be the result of the within Chrysomelidae. Eumolpinae for example is a small proportion of polyphagous Chrysomelidae. Of large subfamily and is represented in our database by Chrysomelidae (including Bruchidae) in Central Europe only one species. Although JOLIVET (2001) attributed only 19 % are polyphagous, whereas monophagous spe- this to the endogenic life of the larvae of Eumolpinae, cies contribute 21 % and oligophagous species 60 % this can only be part of the explanation. KlMOTO (1988) (Schöller 1996). showed remarkable differences in the geographical representation of subfamilies in Chrysomelidae. The trans- Although we know of some invasive species that cause located species originate from all biogeographical re- a lot of harm, for example Leptinotarsa decemlineata gions (Palearctic [47 %], Neotropical [20 %], Nearctic and Diabrotica species, these species had effect on [16 %], Australasian [12 %] and Afrotropical Region [2 plants that were non-native culture plants. The spread of 184 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;Bonner zoologische www.zoologicalbulletin.de;Beiträge 54 (2005) www.biologiezentrum.at

the potato over the world was followed by the spread of new environment, mostly in absence of natural enemies the insect species associated to it. The same holds for or diseases and mostly without competitors. Under more Diabrotica virgifera and the increasing culture of corn natural conditions the herbivorous beetles find less fa- (Zea mais) in Europe. The associated insects will follow vourable conditions. When a leaf beetle enters an eco- the crop when environmental conditions are favourable system it faces a complex of interrelations in which it for the leaf beetle species. Dicladispa armígera (Oliv- first has to establish. It has been shown (KOVALEV ier) (Hispinae) was recorded as a pest of rice in Bengal 2004) in model studies that this process is more likely to and Bangladesh in 1906. It is the most important hispid occur in situations where host plants are evenly spread. pest of rice in tropical Asia, and known from Bangla- Under natural conditions this situation rarely occurs. desh, Myanmar, India, Nepal and China, frequently ELTON (2000) promoted in his famous work on ecologi- causing extensive losses of rice crops. Although D. ar- cal invasions, ecological stability to resist invading spe- mígera is widely distributed throughout India, and fre- cies. quently causes considerable damage to rice crops, it is The threats caused by introduced species on native spe- only considered as a major pest of rice in Andhra cies are larger than can be concluded from this study on Pradesh, Assam, Bihar, Madhya Pradesh, Manipur, Chrysomelidae. Introduced plant species, introduced Kerala, Orissa, Indian Punjab, Uttar Pradesh and West carnivorous species, introduced plant or animal dis- Bengal. It is only a rice pest in the southern provinces of eases, may all have their effect on native leaf beetles. China and is present in western Iran and Sumatra, Indo- Many cases have been listed in which non-native insects nesia and Taiwan (CAB-CPC 2003). (predators or parasitoids) have been released to control Introduced leaf beetle species affecting native plants non-native pest species (for examples release of Oo- seem to be rare. The introduced Plagiodera versicolora, myzus gallerucae Fonscolombe (Hymenoptera, Eulo- Agelastica alni and Xanthogaleruca ¡uteola are known phidae) in the United States for the control of Xan- to damage ornamental trees, and they could cause dam- thogaleruca luteola (Puttler & Bailey 2003). The age to indigenous woody trees such as Salix, Populus, damage of intentionally introduced species to non- Alnus or Ulmus in North America. However this has target, native species from these biological controls are never been reported. At present, Epithrix hirtipennis na- rarely documented. tive to North and Central America occurs in Italy on No negative effects on biodiversity of translocated several Solanaceae, also on the wild and indigenous Chrysomelidae have, at present, been reported. The suc- species (Maurizio Biondi, UAquila-Coppito, pers. comm. cessful control of imported weeds by introduced Chry- 2004) but there is no indication that this causes a threat somelidae can be seen as a positive effect to biodiver- to biodiversity in Italy. sity since some of these weeds prevent natural processes

It seems that negative effects of introduced species on in ecosystems. It is generally accepted that only a mi- nature are barely recognised. However, this does not nority of alien species become abundant. WILLIAMSON mean that they may never occur (DOWNIE 2001). & Fitter (1996) concluded that as many as 80 - 90 % Silfverberg (1995) listed the exotic beetle species that of the established non-indigenous species may actually have colonized Finland and established themselves out- have minimal detectable effects. Apart from that, intro- doors. No leaf beetle was included among the 14 spe- duction of non-indigenous species as a biological con- cies. This might be due to the climatic conditions in trol agent is preceded by intensive research to predict

Finland. However, among the more than 200 invasive the effect of these introductions. BYERS et al. (2002), species listed in the internet invasive species database however, indicated that although there has been a lot of no Chrysomelidae have been listed (IUCN/SSC 2004). research, we still have little information on the effects This database focuses on invasive species that are be- on the communities and species we are attempting to lieved to be a threat to biodiversity. From the absence of protect. They propose key research questions to effec- leaf beetles we might conclude that translocated leaf tively prioritise and manage non-indigenous species. beetles do not threaten biodiversity yet. This is interest- Although there are no cases known in which biodiver- ing because there are very alarming situations of leaf sity is threatened by translocated Chrysomelidae this is beetles attacking cultivated plants (Diabrotica virgifera, no guarantee that it will not happen. Accidental translo- Leptinotarsa decemlineata) or species that have been ef- cations must be avoided as much as possible. No inten- fectively used as control agent against introduced weeds tional translocations should be considered until the fac- (Chrysolina hyperici and C. quadrigemind). In these tors, which limit its distribution and abundance in its circumstances, where alien leaf beetle species eradicate native range, have been understood. The approach sug- a population of a host plant, the host plants grow vigor- gested by the IUCN (1987, 2000) will minimise the ously because of two different situations: one is the cul- risks. tivated situation in which the environmental (fertilisa- tion, hydration) and ecological conditions (absence of The Convention on Biological Diversity (Rio de Janeiro competition, diseases) favour the host plant very much. 1992) called for prevention of the introduction of, con-

A monoculture of good growing plants is the result. The trol or eradication of those alien species, which threaten other is a situation in which an alien plant is invading a ecosystems, habitats or species. However it proved to be © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Ron Beenen: Translocation in Leaf Beetles 185

difficult to discriminate between alien species that are (1996) and Fritzlar (1998) estimate that about 5% of likely to be invasive and those that are not. Because post the leaf beetles of the German states Rhineland and

entry control of invasive alien species is much more Thiiringia came along with introduced plants (neophytes

costly than prevention of invasions, MACK et al. (2000) and archeophytes). suggested that national and international quarantine laws In America conditions were different when transatlantic should be altered by adopting a "guilty until proven in- trade started. In northeastern North America the tribes nocent" approach. were more or less migratory and had hardly progressed

A rarely studied risk of alien species is the role of evo- beyond the Neolithic stage. Therefore in North Amer- lutionary processes. Kovalev (2002) showed changes ica, the chance of native steppe plants invading perma- in the introduced Zygogramma suturalis which devel- nent arable land and transforming into constant weeds oped flight ability and morphological differences within was considerably less than in the old world. When the five generations. The differences where large enough to first Europeans arrived and permanent settlements were attribute the Palearctic population to a new subspecies: founded upon which agriculture started, there were few Z. suturalis volutes KOVALEV. About the risk of hy- indigenous plants present able to intrude as weeds. This bridisation between native leaf beetle species and alien gave free entrance to the corresponding floral elements

species nothing is clear. Hybridisations between native from Europe. Species introduced from Europe into and alien plants species or between genetically different North America are about ten times as numerous as those populations of alien plants are believed to promote rapid transported in the opposite direction (LlNDROTH 1957). evolution and further invasion as has been observed in This is partly explained by the peculiar character of bal-

Spartina anglica (polyploid hybrid of S. alterniflora last traffic in the 19th century. The small sailing vessels from eastern America and the European S. maritima) used for commercial traffic needed cargo or ballast to be and Rhododendron ponticum (introduced from several sailed efficiently. On their way to Northern American different Iberian populations) (PETIT 2004). ports they often were short in tonnage. The bottom of the lower holds were then filled with material available Apart from natural processes as the post glacial spread at the shore of departure: gravel, rocks or even moist of many species (including spread across barriers as the sand. On arrival at their destination the ballast was Alps and Pyrenees in Europe), human-induced translo- dumped on the coast, including the plants and animals cations may have started as soon as man appeared. that came along with this material. Examples of leaf beetles that In Europe the invasion of alien species started in ancient have been transported in this way are Cassida times. The original landscape most probably was a mo- flaveola, Chrysolina staphylea and Gastro- saic of forest and open areas. When man arrived large physa polygoni (LlNDROTH 1957). areas were cleared for agriculture. Plant and animal spe- Not all translocations have to be human-induced. cies from deserts and steppes invaded the European re- Clarke & Zalucki (2004) suggested, based on histori- gion when new, cultivated, habitats appeared. Already cal information, that a substantial population of the at the time of Columbus the coastland of Western monarch butterfly (Danaus plexippus (L.), Danaidae) Europe was almost devoid of forests and transformed was carried to Australia on winds associated with cy- into an artificial steppe inhabited by man's constant fol- clones that hit the Queensland coast in early 1870. The lowers among plants and animals, invited or self-invited leaf beetle Chaetocnema confinis, originally a North (LlNDROTH 1957). Part of theses species have simply American species, has spread to tropical America, Af- expanded their range but it is very likely that part of rica, Asia and the Pacific, accidentally introduced in them were actively transported with seeds or with the Hawaii and established, it is reported to be transported transport of livestock (both plants and animals). It is by a hurricane (JOLIVET 2001). Stegnaspa trimeni is very likely that leaf beetle species associated with crops common on Tristan da Cunha and originated from or the herbs that grow between them have taken the Southern Africa, most probably it was accidentally in- same route as herbs associated with cereals. They are troduced or transported eolic (hurricane ?) (JOLIVET supposed to have arrived in Europe from the Near East 1998). Epithrix hirtipennis from North and Central (PlNHASI et al. 2005). This process is still going on. America was recorded in 1984 from the Azores and It- Hammond (1974) assumed that some of the relative re- aly (DOberl 1994a). Subsequently it spreaded to cent immigrants to the British Isles may represent de- Greece (1988) and Turkey (1993). Although introduc- layed movements of the same kind. An example of such tion with cultivated plants is possible, spreading with a species combination may be Buglossoides (= Lithos- passatwinds is also suggested as a possibility (DOberl permum) arvensis and Longitarsus fitscoaeneaiis RED- 1994a). tenbacher. Both species are nowadays indigenous in

Europe. It is believed that B. a>-\'ensis originated in Acknowledgments. S. M. Clark, M. Biondi. D. Erber (t) southwest-Asia and spreaded with agriculture to large and M. A. Hielkema kindly supported this study either be parts of the temperate parts of the Northern Hemisphere. supplying information or supplying specimens from which In The Netherlands it already occurred during the Ro- conclusions were drawn. Theresa Canning (CAB Interna- man times (WEEDA et al. 1988). It is likely that L. fus- tional) kindly offered a free trial of the internet version of coaeneus spreaded with this herb. FRITZ-KÖHLER the Crop Protection Compendium. P. Jolivet stimulated this 186 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;Bonner zoologische www.zoologicalbulletin.de;Beiträge 54 (2005) www.biologiezentrum.at

study by many references to useful publications. D. G. Downie, R. (ed.) 2001. Alien species friends or foes? The Furth, T. Wagner and an anonymous reviewer made valu- Glasgow Naturalist 23, Supplement: 1-113. able comments on a previous version of this article. Elton, C. S. 2000. The ecology of invasions by animals Participating in the 6th International Symposium on the and plants., With a new forword by D. Simberloff Chrysomelidae was possible with a grant from the University of Chiacago Press, i-xiv, 1-181. Originally published in "Uyttenboogaart-Eliasen Stichting tot bevordering van de 1958 by Methuen, London. entomologische wetenschap". Fields, P. G, Arnason, J. T. & Philogéne, B. J. R. 1988. Distribution of Chrysolina spp. in Eastern Ontario, 18 years after their initial release. The Canadian Ento- REFERENCES mologist 120: 937-938. Fräser, B. & Emberson, R. 1987. Rediscovery of Chry- Allard, M. E. 1 890. Troisiéme note sur les Galérucides. solina quadrigemina (Suffrian) in New Zealand. New Compte rendu des séances de la Société ento- Zealand Entomologist 9: 57-59. mologique de Belgique 4(6): 80-94. Fritz-Köhler, W. 1996. Blatt- und Rüsselkäfer an Ack- Baker, C. R. B., Blackman, R. L. & Claridge, M. F. erunkräutern. Ökologie und Biogeographie in Mittel- 1972. Studies on Haltica carduorum Guerin and aliien europa und Untersuchungen an ungespritzten Acker- beetle released in Britain as a contribution to the bio- randstreifen. Agrarökologie 19: 1-138. logical control of creeping thistle, Cirsium ai~vense Fritzlar, F. 1998. Neozoen unter den Blattkäfern Thürin- (L.) Scop. Journal of Applied Ecology 9: 819-830. gens. Mitteilungen des Thüringer Entomologenver- Beenen, R. 1998. Patterns in the distribution of Galeruci- bandes 5: 61-63. nae in the Netherlands. Museo regionale di Scienze Gonzáles-Megías, A. & GÓMEZ, J. M. 2003. Conse- Naturali -Torino 1998: 7-16. quences of removing a keystone herbivore for the Blossey, B., Cassagrande, R., Tewksbury, L., Landis, abundance and diversity of arthropods associated with D. A., Wiedenmann, R. N. & Ellis, D. R. 2001. Non- a cruciferous shrub. Ecological Entomology 28: 299- target feeding of leaf beetles introduced to control 308. Purple Loosestrife (Lythrum salicaria L.). Natural Ar- Gray, M. E. & Steffey, K. L. 1999. Western Corn Root- eas Journal 21: 368-377. worm adapt to crop rotation. Agrichemical and Envi- Byers, J. E., Reichard, S., Randall, J. M., Parker, I. ronmental News 156: 12-14. M, Smith, C. S., Lonsdale, W. M, Atkinson, L A. Hammond, P. M. 1974. Changes in the British Coleopter-

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naz. ital. Ent., Palermo. Erice, Bagheria: 285-292. SCHÖLLER, M. 1996. Ökologie mitteleuropäischer Author's address: Ron BEENEN, Martinus Nijhoffhove Blattkäfer, Samenkäfer und Breitrüssler. Die Käfer 51, NL - 3437 ZP Nieuwegein, The Netherlands. von Vorarlberg und Liechtenstein 11: 1-65. E-mail: [email protected] 188 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;Bonner zoologische www.zoologicalbulletin.de;Beiträge 54 (2005) www.biologiezentrum.at

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