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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
ESENIAS Training Course, Sofia, 3-4 April 2017
I- Invasion rate: plateau or not plateau ?
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 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
(Seebens et al., Nat.Comm. 2017)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
(Seebens et al., Nat.Comm. 2017)
II- Relationships with trade
ESENIAS Training Course, Sofia, 3-4 April 2017
Clear relationships with trade increase
Values of annually imported commodities
vs. first record rates for all taxonomic groups
(Seebens et al., Nat.Comm. 2017)
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
III- Pathways and donor areas are changing with time
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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 !
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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.)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
(Eschen, Roques & Santini, 2014, Div & Dist.)
ESENIAS Training Course, Sofia, 3-4 April 2017
New trends in colonization of native angiosperms by alien arthropods
Increase in: • Fraxinus • Salix • Castanea Stable in: • Quercus
ESENIAS Training Course, Sofia, 3-4 April 2017
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.)
ESENIAS Training Course, Sofia, 3-4 April 2017
New trends in colonization of native gymnosperms by alien arthropods
Increase in: • Pinus Decrease in: • All others
ESENIAS Training Course, Sofia, 3-4 April 2017
Temporal changes in origin of the established species
New couples: Origin x Donor (imported) tree species
Asia turned dominant as area of primary origin
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
IV- Recently established alien species are spreading faster than before in Europe
Why ?
Arrivals mostly through continental Italy and France
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
Accidentally-introduced species spread faster than intentionnaly- introduced ones
ESENIAS Training Course, Sofia, 3-4 April 2017
No differences in spread speed with regime
ESENIAS Training Course, Sofia, 3-4 April 2017
(Roques et al., 2016 Biol. Inv)
Since 1989, a much faster spread across Europe of alien insects after the establishment
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
The invasive pathway: the transsiberian railway
(Baranchikov, 2013)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
(Baranchikov, Kurteev, 2012) (Orlova- Bienkowskaja, 2013)
2013
Rapid expansion from Moscow towards West
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
V- Genetic tracking of the origin (donor area) is essential
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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%).
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
(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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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 ?
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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, …
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 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 !
ESENIAS Training Course, Sofia, 3-4 April 2017
(Kirichenko et al., Plos One 2017)
VII- How to forecast the next arrivals ?
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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”
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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 ?
ESENIAS Training Course, Sofia, 3-4 April 2017
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 ?
(Eschen, Roques & Santini, 2014, Div & Dist.)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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 ?
ESENIAS Training Course, Sofia, 3-4 April 2017
IX- Climate change proved to promote invasion:
The example of pine processionary moth
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
The example of invasion of Paris Basin
« Invasive » colonies recently detected largely beyond the natural front
(Roques et al., 2014)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
The northwards/upwards shift of isotherms allows survival where it was impossible before
ESENIAS Training Course, Sofia, 3-4 April 2017
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)
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
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
ESENIAS Training Course, Sofia, 3-4 April 2017
Thank your for your attention !