A ROQUES INRA Zoologie Forestière Orléans, France · Roques, 2010 Biorisks . Basic data: EU DAISIE 2005- 2008 . Update: EU JCR- EASIN 2014 . . Rate increase due to phytophages whilst

A ROQUES INRA Zoologie Forestière Orléans, France

PERMIT

GLOBAL WARNING

ESENIAS Training Course, Sofia, 3-4 April 2017

Outlines of the talk

Invasion rate: plateau or not plateau ?

Relationships with trade

Pathways and donor areas are changing with time; a focus on alien insects in Europe

Recently established alien species are spreading faster than before in Europe. Why ?

Genetic tracking the origin (donor area) is essential

Complex patterns of invasion and new genetic admixtures

How to forecast the next arrivals ?

Effect of climate change probable but not easy to ascertain

Climate change proved to promote invasion: The example of pine processionary moth in Europe


I- Invasion rate: plateau or not plateau ?


Recent progresses in invasion studies A novel, worldwide database of established alien species,

including data on first records per country/ continent

45,813 first records for 16,926 species From Plants to mammals

(Seebens et al., Nat.Comm. 2017)


No saturation at world level in the establishment rate of alien species

Global temporal trends in first record rates for all animal and vegetal species at world level



The same trend for most groups but inter-taxonomic variations…

(Seebens et al., Nat.Comm. 2017) ESENIAS Training Course, Sofia, 3-4 April 2017

… as well as inter-continental variations

can be largely attributed to the diaspora of European settlers in the 19th century and to the acceleration in trade in the 20th century



II- Relationships with trade


Clear relationships with trade increase

Values of annually imported commodities

vs. first record rates for all taxonomic groups



Focus on terrestrial invertebrates in Europe Globalization is exponentially accelerating

alien establishment

Roques, 2010 Biorisks

Basic data: EU DAISIE 2005- 2008 Update: EU JCR- EASIN 2014 www.easin.org Rate increase due to phytophages whilst the other groups decreased during the last period ca. 11.5 new phytophagous species per year since the 2000s (4.5 during 1950-75)


Europe

(Roques et al., Biol. Inv. 2016)

Establishment rate is increasing significantly more in species related to woody plants

than in species linked to herbaceous and crops


2,1x more

2-fold more establishments from 2000 on of alien arthropods associated to woody plants

than during 1950- 1975

465 exotic arthropods related to woody plants established in

Europe by 2016

• 405 insects • 60 mites

ca. 8 new species per year

since the 2000s

Likely related to the recent ‘blooming’ in ornamental trade

(Roques, 2010 NZJF updated)


III- Pathways and donor areas are changing with time


Contaminant Deliberate Escape Stowaway

Plants 17% 51% 20% 13%

Mammals 64% 36% 0% 0%

Birds 88% 12% 0% 0%

Amphibia 70% 21% 5% 4%

Marine 7% 30% 30% 33%

Freshwater 59% 31% 7% 3%

Invertebrates 7% 3% 72% 18%

Accidental introductions largely dominant vs. intentionnal ones in invertebrates


Identify a pathway: not so easy !

• Intercepted vs established 1995- 2012: only 7 species over 117 established !

o Anoplophora chinensis (Coleoptera: Cerambycidae) P4P, bonsais o Anoplophora glabripennis (Coleoptera: Cerambycidae), WPM, bonsais o Cydalima perspectalis (Lepidoptera: Pyralidae), P4P o Horidiplosis ficifolii (Diptera: Cecidomyiidae), P4P o Paysandisia archon (Lepidoptera: Castniidae), P4P o Singhella citrifolii (Hemiptera: Aleyrodidae), P4P o Leptoglossus occidentalis (Hemiptera: Coreidae), Hitchhiker (Eschen, Roques & Santini, Div & Dist., 2014)


An example: Most likely pathway for alien insects related to woody plants

Ornamental trade more important than forest products

Plants for planting

Seeds/ fruits

Fresh wood

Wood packaging

0 20 40 60 80 100

% exotic invertebrate species related to woody plants in Europe

437

48

45

17

80

0

3

0

Europe- TotalUK

Europe- Daisie UK- Smith et al., 2007

But based on species’ biology… Actual pathway could be largely different e.g. hitchhiking for a number of species !


The trend tended to increase during the recent period

185

0-18

99

190

0-19

49

195

0-19

74

197

5-19

99

200

0-20

07

048

121620

BarkSeeds/ FruitsWood PackagingRounwood/ Sawnwood

708090

100

Plants for planting

Source: Daisie, 2008

Perc

enta

ge o

f arr

ivals

per

path

way


Another way to identify the invasive pathways ?

Looking at the colonization by exotic insects of tree species growing in Europe

Exotic trees planted in Europe more colonized than native trees The top 10 spp. includes Citrus, Palms, Eucalypts A number of insects arrived along with the exotic host Most did not switch (yet) onto natives (51%)

(Eschen, Roques & Santini, 2014, Div & Dist.)


Looking at large taxonomic

groups during the last period

Faster increase in deciduous,

shrubs, palms

vs.

decrease in conifers and

slowering in fruit trees

Temporal changes in colonization tends to reflect the invasive processes



New trends in colonization of native angiosperms by alien arthropods

Increase in: • Fraxinus • Salix • Castanea Stable in: • Quercus


per family

Large differences between families Fast increase in Myrtaceae (Eucalypts), Fabaceae (exotic legume trees), Rutaceae (Citrus), Arecaceae (palms) but decrease in Pinaceae and Fagaceae

A link with climate change ? Most used as plants for planting and not for wood

(Eschen, Roques & Santini, 2014,

Div & Dist.)


New trends in colonization of native gymnosperms by alien arthropods

Increase in: • Pinus Decrease in: • All others


Temporal changes in origin of the established species

New couples: Origin x Donor (imported) tree species

Asia turned dominant as area of primary origin


Coherent with the temporal changes in the trade of plants for planting

Origin of the import of live plants in the EU Eschen et al., 2014 FEM Asia

NA

AUS- NZ


Plants for planting but not precluding the trade of wood and derivates as a pathway!

Wrong to consider only this pathway with regard to recent invasions:

o Wood packaging still as issue:

Aromia bungii

New outbreaks of A. glabripennis

o Wood trade, too:

exotic ambrosia beetles trapped in Italian ports (Ambrosiodmus rubricollis, Cyrtogenius luteus, Xylosandrus crassiusculus; Rassati et al., 2013)

o Hitchhikers in commodities:

Leptoglossus bugs


IV- Recently established alien species are spreading faster than before in Europe

Why ?

Arrivals mostly through continental Italy and France


It took a long time for many alien species to colonize the whole Europe

Area colonized in 2015 with regard to the date of first record of each alien insect

(Roques et al., 2016)


Accidentally-introduced species spread faster than intentionnaly- introduced ones


No differences in spread speed with regime


(Roques et al., 2016 Biol. Inv)

Since 1989, a much faster spread across Europe of alien insects after the establishment


Arrival during 1989- 2004

Spread before and after the collapse of the Iron Wall and the EU enlargment

Arrival during 1950-1988 Arrival during 1989- 2004 Arrival during 1950-1988


For a number of important species > 15 years to invade all of Europe

vs. dozens of years before

Explosion of ornamental trade combined with collapse of Berlin Wall and release of EU internal border controls (Roques et al., 2016)


Examples of very fast spread West to East I- the American Honeysuckle leaf gall-midge (first

record 2003)

Honeysuckle leaf gall-midge Obolodiplosis robiniae (North America) 2003

2004

2006

2009

2007

2008

2010

2011


The elm zigzag sawfly (China) Aproceros leucopoda

2003

2006

2009

2013

2010

2011

Examples of very fast spread West to East II- The Asian elm zigzag sawfly (first record 2003)


Faster spread especially obvious for recent aliens associated with eucalypts and palms

Eucalypts: foliage feeders and gall-makers (Ophelimus maskelli, Leptocybe invasa, Glycaspis brimblecombei, Blastopsylla occidentalis, Thaumastocoris peregrinus) colonized most of the Mediterranean countries in less than 10 years

vs.

a much slower colonization rate by Phoracanta spp. (arrival 1969 and 1992) and Gonipterus spp. (arrival 1975 and 1991)


An invasion East to West: Polygraphus proximus, a threat for European fir

moving westwards with new fungal association

Introduced from the Russian Far East to Southern Siberia presumably late1980s Beetle’s success related to lack of fir resistance to an associated blue stain fungus Grosmannia aoshimae

(Baranchikov, 2013)


The invasive pathway: the transsiberian railway

(Baranchikov, 2013)


Another East- West highly-threatening dispersal EAB progression westwards from European Russia

wind protection forest belt (near the town of Puschino, 2012) (Photo: Y. Baranchikov)

Human-aided transport from the native Far East First record in Moscow in 2003


(Baranchikov, Kurteev, 2012) (Orlova- Bienkowskaja, 2013)

2013

Rapid expansion from Moscow towards West


An accelerating effect of the Ukrainian conflict ?

Dark gray: invasive range of A.

planipennis

Light gray: native range of A.

convexicollis

Triangles: localities where A.

convexicollis was found outside its native range in 2007–2013.

Bienkowskaja & Volkovitsh Biol Invasions (2015)

EAB invasion is favoring the expansion of a native species related to ash, A. convexicollis

2013


V- Genetic tracking of the origin (donor area) is essential


Example of very fast spread West to East The Asian box tree moth, Cydalima perspectalis

Probably arrived from China in 2005

2006

2007

2008

2011

2010

2014

2012

2013


Rapid spread towards Central Asia along the Black sea, resulting in heavy damage

A north route with the Sotchi games and a south route from Turkey Krasnodar- 2012 Sotchi- 2013 Chechen Rep- 2013 Abkhazia, Georgia (Adjara, Samegrelo- 2015 but probably 2013)

Heavy damage in Adjara in 2015 (M. Kenis courtesy)


Genetic analysis revealed the probable routes Multiple introductions followed by human-mediated

dispersal with ornamental trade

Spatial genetic structure with 3 geographic regions (West Europe, Central Europe, East Europe and Central Asia)

Bras et al, 2016

COI, mtDNA


Several introduction events followed by a

spread within Europe (Javal et al., 2016)

A similar scenario for the highly-threatening Asia long-horned beetle, Anoplophora glabripennis


Genetic markers proved the horse-chesnut leaf miner, Cameraria ohridella

to originate from the Balkans

Valade et al (2009), Molecular Ecology

Photo: D. Lees

COI mtDNA (barcode fragment - 658 bp)

25 haplotypes Only haplotype “A” is dominant (67–100%).


Confirmed by analysis of herbarium and sequencing of old samples (Lees et al., 2011, Frontiers Ecol. & Envir.)

VI- Complex patterns of invasion and new genetic admixtures


(Lombaert et al., 2011, PLoSOne)

Bridgehead effect A particular invasive population serves as a source for subsequent invasions (Lombaert et al. 2010)

• Genetic recombinations from different sources

• Adaptative patterns and genetic drift

• Hybrids more aggressive

The Harlequin ladybird, Harmonia axyridis


Temporal patterns of invasion by seed bugs, Leptoglossus occidentalis

Western American conifer bug invasion in Europe (Leptoglossus occidentalis) from its arrival in Italy in 1999

2002 2004

2004

2007

2007

2007

2010

2011

2011

2006

2006

2006

2006

2009

2009

2009

2009

2008

2008

2008

2008

1999 2005

2003

2003

2003

Lesieur et al., submitted


DNA analyses and Approximate Bayesian Computation (ABC) approach : Best scenario for the origin of the population of Northern Italy

eNA Italy wNA

Unsampled population in eNA

wNA population admixture eNA-wNA

X

X X

A Western US (bug native range) origin for the European invasion ?


Unsampled population in eNA

X

Admixtures…

DNA analyses and ABC techniques: Best scenario for the origin of the population of Barcelona, Spain

Multiples introductions and bridgehead effect in bug invasion


The most reliable scenarios for Leptoglossus European invasion

Source for all: not wNA but eNA= Bridgehead effect

Multiple independent introductions : - at least 2 in Italy and Spain

Movements within Europe: flight+ hitchhiking

- Probably much more : NW France, Spain, …


Temporal patterns of invasion by the blue gum gall wasp, Leptocybe invasa, in the Mediterranean basin

Direct trade of Eucalypts with Australia more limited: Exchanges of ornamental plants for planting and/or cut foliage likely responsible for the invasion… from one Mediterranean country to the others


Bridgehead effect probably more common than expected in relation with the explosion of ornamental trade

Significant consequences

Important to consider systematically the donor area, not only the native range

Necessity to test for behavioral consequences of potential hybridization from different genetic sources and from populations having been submitted to strong bottlenecks


Even more complex: Where does come from the Lime leaf miner, Phyllonorycter issikii ?

East or West ?

Moscow

- Tilia range - P. issikii invasion

*2009 *2008

*2011

*1985 *1999

*2006

*1992

*1995

*1992 *1995 *2004 *2004 *2005

*2003

*2008

*2010 *2012

*1977

Moscow

(Kirichenko et al., Plos One 2017)


6 unique haplotypes in RFE

1 unique haplotypes in Japan

COI mtDNA (barcode fragment - 658 bp)

Contradictory high genetic diversity in the putative invaded area vs the native one !


(Kirichenko et al., Plos One 2017)

VII- How to forecast the next arrivals ?


Relying on interceptions to predict the arrivals ? Large discrepancies with establishments !

Intercepted vs established 1995- 2012: (Eschen, Roques & Santini, 2014, Div & Dist.)

The major problem: only A1 and A2 pests targeted Most new species: unknown as pests in countries of origin (cf EAB)


How to forecast and detect precociously the next arrivals ? A number of EU projects to develop novel strategies

Preventive warning: Lists based on the pests known in the native range: most

newly-arrived species not considered as pests … or Unknowns Sentinel plantings of European plants in exotic countries Sentinel nurseries of exotic plants in exotic countries Survey of arboreta and botanical gardens with European plants

Former EU projects PRATIQUE, ISEFOR & PERMIT At present COST ‘GLOBAL WARNING’ and IPSN

Early detection at arrivals: Test of multiplex traps and lures with generic attractivity in

ports-of entry French PORTRAP project (2015-2016), proposed EUPHRESCO project

“Multi-lure and multi-trap surveillance for invasive tree pests”


Beijing suburban area Continental conditions INRA- CABI

Fuyang, nr. Hangzhou Warm and humid climate INRA- IOZ- ZAF

Sentinel plantings: 7 European trees species in China (2008-2011) survey for 3 years for insect and pathogen colonization


Experiments designed for statistic analysis of colonization by insects and pathogens

100 seedlings per site and species Random planting of blocks of 25 seedlings

Abies alba Quercus suber Cupressus sempervirens Quercus petraea Fagus sylvatica Quercus ilex Carpinus betulus

….. ….. ….. ….. …..

….. ….. ….. ….. ….. ….. ….. ….. ….. …..

….. ….. ….. ….. …..

….. ….. ….. ….. …. ….. ….. ….. ….. ….. ….. ….. ….. ….. ….. ….. ….. ….. ….. …..

….. ….. ….. ….. ….. ….. ….. ….. ….. …..

….. ….. ….. ….. …..

….. ….. ….. ….. ….

All seedlings individually tagged


105 insect species colonized the seedlings in 3 yrs 3 years enough to check species recruitment Top species not incidental> 15 occurrences over several years: A list of 39 threatening species

5 candidates for PRA

Pteroma nr. pendula (positive tests under

quarantine conditions)

Same for pathogens ESENIAS Training Course, Sofia, 3-4 April 2017

Sentinel nurseries taking into account the role of ornamental trade (2012- 2014) 71 insect species in >20 families, most not included in Chinese lists

Kenis et al., in prep.

Selection of the 6 most imported woody plants from China to Europe during 2008-2011 No treatments to follow the « freely » colonization by insects and pathogens Large differences among tree species after 3 yrs Would have allowed to warn about the arrival of the invasive boxmoth, Cydalima perspectalis


COST GLOBAL WARNING: An attempt to promote sentinel strategies … but there are limits

• A manual in preparation for assigning observed damage to insects and pathogens in broadleaved and conifer sentinel plants (End 2016)

• Many specimens (larvae, eggs) not identifiable by morphological keys (some may be unknown to Science)

• Necessity of molecular librairies (barcodes) but nothing for some taxa • Mostly foliage pests, no pests of mature trees (xylophagous,

seed/ fruit insects, …)

• Role of travel and plantation stress difficult to assess


Arboreta surveys another method for identifying potential invaders

and for early detection of invaders

- Inspection of European trees and shrubs in arboreta and other plantations in other continents

- Inspection of arboreta/ gardens in Europe

IPSN Conference, Kew, UK, 24 February 2016

A significant example : Survey of rose seed infestation by seed chalcids in major European botanical gardens

Survey 2014- 2016: Detection of 5 exotic species of Megastimus seed chalcids in MNHN Paris and Kew gardens

No alien chalcids outside these gardens yet: Arboretum= trap for aliens ?... and early warning Pathway= uncontrolled seed exchange (no X-rays) between gardens ?


VIII- Faciliting effect of climate change probable but not easy to ascertain

Faster increase in colonization of tropical/ subtropical alien trees vs. natives

Link with • Climate change ? • Change in trade because most

exotics used as plants for planting and not for wood ?

• Both ?



The rate of colonization of exotic Eucalypts much faster than that of native deciduous

Since 2000: 2000: Ophelimus maskelli 2002: Ctenarytaina spatulata 2003: Leptocybe invasa 2006: Blastopsylla occidentalis 2006: Ctenarytaina peregrina 2007: Glycaspis brimblecombei 2011: Thaumastocoris peregrinus

A further analysis at the tree species level necessary


The rate of colonization of exotic Citrus much faster than that of native fruit trees

Since 1998: 1998- Brevipalpus californicus 1998- Brevipalpus phoenicis 1998- Chrysodeixis acuta 1998- Singhiella citrifolii 2000- Acaudaleyrodes rachipora 2000- Anoplophora chinensis 2001- Eutetranychus banksi 2001- Icerya formicarum 2007- Acharia stimulea 2007- Aleuroclava aucubae 2008- Aleurocanthus spiniferus 2009- Lopholeucaspis japonica


The same story for alien insects on palms

Palm weevil, Rhynchophorus ferrugineus, palm moth, Paysandisia archon, leaf beetles, … The invasive moth and weevil recorded from the Canary islands or continental Spain from the late 1990s… but without spreading elsewhere 2004- 2007: Colonization of the whole southern Europe One certainty: the role of Athens’ Olympic games in 2004 with large imports of Spanish palms Warming up: release of thermal barriers for expansion ?


IX- Climate change proved to promote invasion:

The example of pine processionary moth


Pine processionary moth (PPM), a model for climate change (IPCC)

5 larval instars

Possible prolonged diapause

An insect native of the Mediterranean Basin, with a winter larval development


TEMPERATURE

>-16°C

LONG-TERM SURVIVAL

(Buffo et al. 2007)

Realized Feeding Threshold (RFT)

T nest > 9°C (day)

Activation T

T air > 0°C (following night)

Potential feeding T

+

FEEDING

(Battisti et al. 2005)

IMMEDIATE SURVIVAL

Winter warming up= release of thermal constraints constraining PPM range


A latitudinal expansion northwards (and in altitude)

A 100 km latitudinal shift in the Paris basin since 1972 with a significant acceleration since 1992

Robinet et al. 2010

Unpubl. data

Northwards shift - by 2.6 km/yr from 1972 to 2011 - by 5.5 km/yr from 1996 to 2011

whilst mean winter temperature increased by 1°C

1971 1992 1996 2005 2011

Mean minimum temperature from October to March in Orléans (moving mean 5 yrs)

2.46°C +/- 0.86°C (SD)

3.55°C+/-0.68 (SD)

1

2

3

4

1972 1990 1980 2000


Map of PPM expansion all over Southern Europe The 2012 front edge

PCLIM Project INRA, with the help of A. Battisti (Italy) P. Mirchev (Bulgaria) G. Georgiev (Bulgaria) M. Dautbasic (Bosnia) D. Matosevic (Croatia) K. Ipekdhal (Turkey) M. Glavendekic (Serbia) I. Papazova (Macedonia) S. Naceski (Macedonia) D. Avtzis (Greece) E. Cota (Albania) T. Manole (Romania) J. Garcia (France, Switzerland) F. Goussard (France, Switzerland)

(Roques et al, 2015)


Only expanding ? No ! Also a man-mediated invader …. although seems crazy to translocate an urticating pest !

A number of colonies appeared far beyond the expansion front since the early 2000s Were do they come from ? The expanding area or far beyond ? How can they survive far beyond the natural front ?

2008- 190km 2012- 90km

2003- 2008- 14-60km

2015


The example of invasion of Paris Basin

« Invasive » colonies recently detected largely beyond the natural front

(Roques et al., 2014)


Limited female flight capabilities and particular population genetic structure Colonies = long-distance jumps from southern areas !

Females fly at more 11km (a few)

Robinet et al. Biol. Inv. 2012

DNA microsatellites assigned colonies to southern populations, not to front


Convergent genetic and biological data: Populations not coming from the front edge but from very long distances

The likely pathway: The trade of mature trees moving processionary pupae with soil from southern France (> 800 km) (Robinet et al.,

2012 Biol Inv)

The same pathway for the invasion of UK by the oak processionary moth (Evans,

2008)

Many PPM nests


The northwards/upwards shift of isotherms allows survival where it was impossible before


0.05

0.1

0.15

0.2

0.25

0.3

0.35

0.4

0.45

0.5

6.2 6.4 6.6 6.8 7 7.2 7.4 7.6 7.8 8 8.247.6

47.8

48

48.2

48.4

48.6

48.8

49

49.2

période 1988-1997 période 1998-2007

Obernai Strasbourg

Metz Saarebrücken

Nancy

Basel

Karlsruhe

6.2 6.4 6.6 6.8 7 7.2 7.4 7.6 7.8 8 8.247.6

47.8

48

48.2

48.4

48.6

48.8

49

49.2

Establishment impossible in the early 1990s; colonies established since 2007 from accidental transportation with black pines used for runabouts

Obernai (French- German border):

Most invading colonies proceed from management using ornamental plantations of mature pines

Ornamental pines along as corridors and relays for PPM dispersal (Rossi et al., 2015)


Urban heated islands: PPM survival is even favoured in large cities once colonized

1972

1992

1996

2004

2009

PARIS

Orléans

Pontoise Severe 15-day cold period in the Paris basin during January 2009

-12

-12.5

-13

-14

-13

Absolute minimum T

Nb hours where T<0

79.1%

75.8% 75.9%

81.7%

70.5% 71.6%

75.1% 74.0%

1972

1992

1996

2004

2009

PARIS

Orléans

Pontoise

% larval survival

Robinet et al. Biol. Inv. 2011


A changing parasitoid complex: Expanding PPM populations escape from egg parasitoids

Colonies beyond front with only pupal parasitoids

a a a

a a a a ab

b b

bc

b

c

c

c

bc hole of caterpillar

hole of parasitoid

Significant decrease in parasitism from endemic areas to front edge

Less egg mortality may allow larger PPM colonies at the front edge and in isolated colonies, ie better winter survival in colder places

(Imbert, 2012)

No parasitoids or only pupal ones

High frequency of egg parasitoids


All of Europe and Minor Asia likely to become favorable for PPM invasion

in the near future

(Robinet et al., 2015)

Any long-range transport with potted pines from the native range is likely to generate colony establishment

Probability of PPM establishment under different temperature increase scenarios


An important question remains: PPM adaptations at the front edge and in colonies beyond front ?

Different genetic markers show that the front population is a mosaic of

new, unexpected genetic combinations

Possible adaptations at the front edge - Larger flight capabilities and adapted

morphology and carbohydrate content ?

- Better cold resistance ?

- Advanced phenology ?

- Fertility (Egg load)

- Nest size

- Lower urticating capabilities (no. setae and allergen content) in relation to escape from natural enemies

- Increased man/ animal sensitivity in newly-colonized zones ?

Paris Basin 2015- 2016


Invasions are accelerating with globalization without any saturation

Insect invaders mostly arrive with plants for planting but wood and derivates are still to be considered

Hitchhiking is a serious problem difficult to be adressed

Present EU inspections simply focusing on quarantine pests are not efficient

New combinations origin * commodity have to be focused on

Possible bridgehead effects have to be systematically considered as well as the donor area and not only the native area of the species

Possible new associations (insect* fungus*nematode) between invaders and natives to be systematically checked

The release of barriers in EU accelerate the invasive spread

Relationships between invasions and climate change are still a black box in most cases although recent establishement of species related to subtropical hosts suggest a facilitation.

Such facilitation is proved for the pine processionay moth whose actual expansion results from the combination of global warming, the development of ornamental pine plantations, and the increasing trade of mature trees

Conclusions and take- home messages


Thank your for your attention !

Documents

A ROQUES INRA Zoologie Forestière Orléans, France · Roques, 2010 Biorisks . Basic data: EU DAISIE 2005- 2008 . Update: EU JCR- EASIN 2014 . . Rate increase due to phytophages whilst