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CLIMEX models for Tamarixia triozae Logan DP and Gardner-Gee R November 2012 A report prepared for New Zealand Tamarillo Growers Association Inc. DP Logan Plant & Food Research, Te Puke R Gardner-Gee Plant & Food Research, Mt Albert SPTS No. 7671

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Page 1: CLIMEX models for Tamarixia triozae · 2019. 4. 6. · CLIMEX models for Tamarixia triozae. SPTS No. 7671 Executive summary CLIMEX models for Tamarixia triozae David Logan and Robin

CLIMEX models for Tamarixia triozae

Logan DP and Gardner-Gee R

November 2012

A report prepared for

New Zealand Tamarillo Growers Association Inc.

DP Logan

Plant & Food Research, Te Puke

R Gardner-Gee

Plant & Food Research, Mt Albert

SPTS No. 7671

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DISCLAIMER

Unless agreed otherwise, The New Zealand Institute for Plant & Food Research Limited does not give any prediction, warranty or assurance in relation to the accuracy of or fitness for any particular use or application of, any information or scientific or other result contained in this report. Neither Plant & Food Research nor any of its employees shall be liable for any cost (including legal costs), claim, liability, loss, damage, injury or the like, which may be suffered or incurred as a direct or indirect result of the reliance by any person on any information contained in this report.

LIMITED PROTECTION

This report may be reproduced in full, but not in part, without prior consent of the author or of the Chief Executive Officer, The New Zealand Institute for Plant & Food Research Ltd, Private Bag 92169, Victoria Street West, Auckland 1142, New Zealand.

PUBLICATION DATA

Logan DP, Gardner-Gee R. 2012. CLIMEX models for Tamarixia triozae. A report prepared for: New Zealand Tamarillo Growers Association Inc.. Plant & Food Research milestone: 49962, Job Code. P/333005/01. SPTS No. 7671..

This report has been prepared by The New Zealand Institute for Plant & Food Research Limited (Plant & Food Research), which has its Head Office at 120 Mt Albert Rd, Mt Albert, Auckland.

This report has been approved by:

David Logan

Scientist, Applied Entomology

Date: November 2012

Louise Malone

Science Group Leader, Applied Entomology

Date: November 2012

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Contents

Executive summary i 

1  Introduction 1 

2  Method 2 2.1  General methods 2 2.2  Selection of parameter values 2 

3  Interpretation of CLIMEX outputs 6 3.1  Identify areas in New Zealand where T. triozae is likely to do

particularly well 6 3.2  Identify areas in New Zealand where T. triozae is unlikely to establish 6 3.3  Examine how well T. triozae is likely to establish in Bay of Plenty,

Northland, and coastal Taranaki 6 3.4  Examine how well T. triozae is likely to establish in Pukekohe,

Hawke’s Bay, Manawatu and Canterbury 6 3.5  Examine the climate match between California and Hawke’s Bay 6 

4  Summary 8 

5  Acknowledgements 31 

6  References 31 

Appendix Table A1. Collection sites for Tamarixia triozae in North and Central America. 33 

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The New Zealand Institute for Plant & Food Research Limited (2012) Page i CLIMEX models for Tamarixia triozae. SPTS No. 7671

Executive summary CLIMEX models for Tamarixia triozae

David Logan and Robin Gardner-Gee, November 2012, SPTS No. 7671

This report details work undertaken for PFR contract no.28844, for New Zealand Tamarillo Growers Association, Potatoes NZ and Heinz Watties NZ Ltd for the period 1 July 2012 to 31 October 2012. The climate modelling tool CLIMEX was used to assess the suitability of the New Zealand environment for the survival and persistence of Tamarixia triozae (Burks) (Hymenoptera: Eulophidae), a potential biological control agent for the tomato potato psyllid Bactericera cockerelli (Sulc) (Hemiptera: Triozidae).

CLIMEX models of T. triozae were developed using published and unpublished records of distribution and on currently known biology. Ecoclimatic Index scores, the standard CLIMEX model output, were mapped in the GIS software ArcMap 10.0. As biological information on the response of T. triozae to moisture and temperature is minimal, multiple models were developed to fit the known distributions in Central and North America. The models simulate two alternative responses to temperature and three alternative responses to moisture. The models may be characterised as:

Model 1 (warm-dry preference, cold and wet limited)

Model 2 (hot-dry preference, cold and wet limited)

Model 3 (warm-dry preference, wet limited)

Model 4 (hot-dry preference, wet limited)

Model 5 (warm-dry preference, wet tolerant)

Model 6 (hot-dry preference, wet tolerant).

All six models indicate that the east coast of the South Island is probably suitable for T. triozae. Suitability of other areas is dependent on whether T. triozae is more moisture tolerant than models 1–4 assume. Areas of the east coast of the North Island, particularly Hawke’s Bay, and areas around Auckland, Waikato and the Manawatu/Wanganui may be suitable when the lack of tolerance of T. triozae for moist conditions is relaxed (models 5 and 6, wet tolerant). Some localities may also be suitable for T. triozae in Northland and the Bay of Plenty.

As tomato potato psyllid has a similar range in North and Central America to T. triozae and has established successfully in New Zealand, it is probable that T. triozae can also establish in New Zealand. Caution is needed when interpreting the mapped outputs of the CLIMEX models. Maps of suitable habitat in New Zealand for T. triozae depend on model assumptions about the moisture and temperature preferences of T. triozae. Uncertainty about T. triozae responses to abiotic variables and even the extent to which abiotic variables determine the range of T. triozae in North and Central America means that maps produced in this report are necessarily speculative.

For further information please contact:

David Logan The New Zealand Institute for Plant & Food Research Limited 412 No. 1 Road, RD2, Te Puke, 3182, New Zealand [email protected] 07 9289794

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The New Zealand Institute for Plant & Food Research Limited (2012) Page ii CLIMEX models for Tamarixia triozae. SPTS No. 7671

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The New Zealand Institute for Plant & Food Research Limited (2012) Page 1 CLIMEX models for Tamarixia triozae. SPTS No. 7671

1 Introduction The eulophid parasitoid Tamarixia triozae (Burks) is being considered for release into the New Zealand environment to control the tomato potato psyllid Bactericera cockerelli (Sulc), a major new pest of solanaceous crops (Figure 1). Here we report on CLIMEX modelling to assess whether the climate of cropping areas in New Zealand may be suitable for T. triozae establishment and persistence. The report follows previous host preference studies to quantify risk to native psyllids of parasitism by T. triozae (Gardner-Gee 2012) (Figure 1).

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2 Method

2.1 General methods

CLIMEX (V2.0) (http://www.hearne.com.au/attachments/ClimexUserGuide3.pdf) was used to simulate the possible distribution of T. triozae in New Zealand. CLIMEX is model-building software to simulate a species’ geographical distribution based on climatic variables (Sutherst & Maywald 1991). A CLIMEX model generates indices that describe population growth and others that describe limits to growth based on five meteorological variables (minimum and maximum temperature, rainfall, and relative humidity at 0900 and 1500 h). Indices that may be used to promote growth are dormancy/chilling, temperature, moisture and light. Limits to growth imposed by adverse climatic conditions are grouped into cold, hot, dry and wet stresses and combinations of two states (cold-wet, cold-dry, hot-wet and hot-dry). The most frequently used reported outputs for CLIMEX models are Ecoclimatic Index (EI) scores. They are a combination of growth indices and stress indices and are scaled between 0 (unsuitable) and 100 (very suitable). EI scores can be interpreted by assuming that there is a favourable and unfavourable season for insect growth (Sutherst 2003). For example, winter in temperate regions is a period when insect populations have limited growth and may enter diapause when no growth occurs. A consequence of this assumption is that most population growth only occurs in part of the year. An EI score of 30 indicates that maximum growth has been achieved in at least 60% of the favourable season. An EI score of <10 indicates suboptimal growth conditions and/or high stress conditions.

Models for T. triozae were constructed in CLIMEX (V2.0) to match its known distribution in North and Central America (Appendix, Table 1). Models were also based on some limited biological data. As there is considerable uncertainty for many model parameters (Table 1), multiple models were generated. Simulations were based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004; Stephens et al. 2007). Mapped output for interpolated climate data (i.e. CX_CRU_61-90_V2) is different from that for weather station data and may lead to differences in interpretation of suitability at local scales. For comparison, some simulations for New Zealand were based on weather station data using the CLIMEX climate database metdata.mm. This database contains records for over 3000 weather stations worldwide. I (DPL) modified by the database by addition of 285 sites for New Zealand and 30 sites to California to increase map resolution. Climate data for additional sites in New Zealand were provided by S. Worner (Lincoln University, New Zealand). Climate data for extra sites in California were from the University of California IPM online site (http://www.ipm.ucdavis.edu/WEATHER/index.html accessed 21 and 22 June 2012). The EI scores for North and Central America and for New Zealand simulations were mapped with GIS software (ArcMap 10.0).

2.2 Selection of parameter values

Collection localities for T. triozae are in arid and semiarid areas in mid-west USA and in central Mexico (Pletch 1947; Appendix Table A1). The distribution of collection localities suggests that T. triozae has a preference for warm-hot and dry environments. Survival in dry environments does not necessarily preclude survival in more moist environments. For example, the tomato potato psyllid shares a similar distribution in North America to T. triozae and likewise may be considered to prefer arid and semiarid environments. However in New Zealand, tomato potato psyllid is abundant in crops at Pukekohe, a wetter area than the US central Midwest and central Mexico (annual rainfall at Pukekohe c. 1100 mm v. US mid-west c. 200-800 mm).

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Collection from sites at high latitudes (45-47o N) suggests that T. triozae may be able to overwinter there and is tolerant of significant chilling (< -5oC). It is possible that T. triozae enters diapause to survive cold periods as occurs in some other eulophids (Nechols et al. 1980; Hamerski et al. 1990).

Parasitism by T. triozae was reported to occur relatively late in the phenology of tomato potato psyllid in Colorado (Johnson 1971). This may indicate that T. triozae has a relatively higher minimum temperature threshold for development than B. cockerelli. The lower developmental threshold temperature (DV0) for B. cockerelli has been estimated as 7.1 for potatoes and 7.5 for tomatoes by Tran et al. (2012). No estimates of DV0 are available for T. triozae, as temperature-dependent development has not yet been modelled. The development period of T. triozae has only been determined at one constant temperature (12 d at 26 ±1oC) (Rojas 2010). Thermal biology of congeneric species can sometimes be similar (Ikemoto 2003) and may be a useful guide where data are not available. Some temperature-dependent development rate data are available for T. leucaenae (Patil et al. 1993), a native of central America with a type locality of Trinidad. Development rate data are also available for T. radiata, which has a wide semiarid to tropical distribution (Gomez-Torres et al. 2012). As the distribution of T. triozae is closer to that of T. leucaenae than T. radiata, data from Rojas (2010) for T. triozae and from Patil et al. (1993) for T. leucaenae were combined to give an estimate of DV0 and thermal constant (day-degrees PDD above DV0 required to complete development from egg to adult). The thermal constant estimated in this way (160 PDD) is less than half that estimated for development of tomato potato psyllid on potatoes (358 PDD) and tomatoes (368 PDD) by Tran et al. (2012).

As biological information on the response of T. triozae to moisture and temperature is minimal, multiple models were developed to fit the known distribution in central and North America. The models simulate two alternative responses to temperature and three alternative responses to moisture. The models may be characterised as:

Model 1 (warm-dry preference, cold and-wet limited)

Model 2 (hot-dry preference, cold and wet limited)

Model 3 (warm-dry preference, wet limited)

Model 4 (hot-dry preference, wet limited)

Model 5 (warm-dry preference, wet tolerant)

Model 6 (hot-dry preference, wet tolerant).

Preferences for warm (models 1, 3, 5) or hot (models 2, 4, 6) temperatures were simulated by varying the lower developmental threshold (DV0), the optimum range for growth (DV1-DV2) and upper limit for population growth (DV3) (Table 1). As models 1, 3 and 5 have a relatively low DV0 and DV1, they simulate growth in more areas of New Zealand than models 2, 4 and 6. Parameters describing cold stress (DTCS, DHCS) were selected to allow some survival in southern Canada. For models 1-4, parameters for moisture range allowing population growth (SM0, SM1, SM2, SM3) were selected to limit the distribution of T. triozae to the drier regions of central USA and to Mexico. In models 5 and 6, the upper moisture range allowing population growth was expanded and wet stress (SWWS, HWS) was relaxed. In this scenario, the absence of T. triozae in the eastern USA may be due to non-climatic variables such as tomato potato psyllid distribution. For some species, combinations of temperature and moisture can interact to limit distribution of a species when either factor alone does not (e.g. Smith 2012). In models 1 and 2, I used the stress combination of wet-cold (DTCW, MTCW, PCW) to limit the distribution

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of T. triozae in eastern USA (Table 1). In models 3 and 4, I used wet stress alone to limit the simulated distribution of T. triozae in wet tropical areas such as Florida, where it does not occur.

Each model was applied to Central and North America and the EI scores generated for interpolated climate data were mapped, together with collection localities, in ArcMap 10.0 (Figures 2-7). Differences between EI scores generated by models 1 and 3 (wet stress v. cold-wet stress) (Figure 8), models 3 and 4 (hot temperature v. warm temperature preference) (Figure 9) and by models 4 and 6 (moisture-intolerant/dry v. moisture-tolerant preference) were mapped to highlight dissimilarities between projected geographic distributions (Figures 8-10). Models were then applied to New Zealand, interpolated (Figures 11-16), and weather station (Figures 17-19) climate data and EI scores mapped. Because maps of EI scores were for the hot preference models 2, 4 and 6, they are more conservative than they would otherwise be.

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Table 1. CLIMEX parameters for models describing potential habitat suitable for Tamarixia triozae. Parameter group Parameter Model 1 Model 2 Model 3 Model 4 Model 5 Model 6

Temperature Index

Lower Temperature Threshold (DV0) Lower Optimum Temperature (DV1) Upper Optimum Temperature (DV2) Upper Temperature Threshold (DV3) Minimum degree-days above DV0 to complete a generation or thermal constant (PDD)

9 16 28 33

160

12 20 28 35

160

9 16 28 33

160

12 20 28 35

160

9 16 28 33

160

12 20 28 35

160

Moisture Index

Lower Soil Moisture Threshold (SM0) Lower Optimal Soil Moisture (SM1) Upper Optimal Soil Moisture (SM2) Upper Soil Moisture Threshold (SM3)

0.05 0.1 0.8 1.0

0.05 0.1 0.8 1.0

0.05 0.1 0.8 1.0

0.05 0.1 0.8 1.0

0.05 0.1 1.0 1.1

0.05 0.1 1.0 1.1

Cold Stress Cold Stress Degree-Day Threshold (DTCS) Cold Stress Degree-Day Rate (DHCS)

-6 -0.0002

-6 -0.0002

-6 -0.0002

-6 -0.0002

-6 -0.0002

-6 -0.0002

Wet Stress Wet Stress Threshold (SMWS) Wet Stress Rate (HWS)

1 0.025

1 0.025

1.2 0.005

1.2 0.005

Cold Wet Stress Cold–Wet Degree-day Threshold (DTCW) Cold-Wet Moisture Threshold (MTCW) Cold-Wet Stress Rate (PCW)

60 1 0.05

60 1 0.05

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3 Interpretation of CLIMEX outputs Five questions were posed for climate modelling of the potential distribution of T. triozae in New Zealand using CLIMEX.

3.1 Identify areas in New Zealand where T. triozae is likely to do particularly well

All six models indicate that the east coast of the South Island is probably suitable for T. triozae (Figures 11-19). Larger areas of the east coast of both North and South Islands and areas around Auckland and Waikato may be suitable when the lack of tolerance of T. triozae for moist conditions is relaxed (models 5 and 6, wet tolerant) (Figure 19).

3.2 Identify areas in New Zealand where T. triozae is unlikely to establish

All models indicate that the west coast of the South Island is probably unsuitable for T. triozae to establish (Figures 11–19). Similarly, the central North Island and parts of the Bay of Plenty, Northland, and Taranaki are probably unsuitable.

3.3 Examine how well T. triozae is likely to establish in Bay of Plenty, Northland, and coastal Taranaki

Based on cold x-wet limited and wet limited models (models 1–4) the Bay of Plenty, Northland, and Taranaki are probably unsuitable for T. triozae. Models 5 and 6 simulating wet tolerance indicate that there may be some localities that are suitable for T. triozae in Northland and the Bay of Plenty (Figure 19).

3.4 Examine how well T. triozae is likely to establish in Pukekohe, Hawke’s Bay, Manawatu and Canterbury

Models 5 and 6 simulating wet tolerance indicate that Hawke’s Bay, Manawatu/Wanganui and Canterbury are suitable for T. triozae. Models 1-4 indicate that areas other than Canterbury are not suitable or relatively less suitable than Canterbury for T. triozae.

3.5 Examine the climate match between California and Hawke’s Bay

California’s climate varies from the hot dry south with annual rainfall of <100 mm and average extreme maximum and minimum temperatures of c. 40°C and 5°C to the milder climate of the north coast with annual rainfall of ≥2000 mm and average extreme maximum and minimum temperatures of c.17°C and 5°C. Subzero temperatures occur in eastern inland areas. Annual rainfall totals in the central valley, the key agricultural area of California vary from 100 mm in parts of Kern County in the south to 750 mm in northern counties. Low rainfall in the main production areas of the San Joaquin Valley, Tulare basin and Imperial Valley means that agricultural production depends heavily on irrigation.

Hawke’s Bay has average extreme maximum and minimum temperatures of 24°C and 2°C with annual rainfall of c. 800–900 mm.

Climate in Hawke’s Bay and California may be compared on the basis of the CLIMEX models for T. triozae (Table 2). In general, EI scores for Hawke’s Bay weather stations are high (>30) and indicate that the climate is closely matched to the estimated climatic preferences of

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T. triozae. Weather stations in the central valley of California (agricultural area) had EI scores that range from relatively poorly matched and probably unsuitable for T. triozae (3–5) to relatively closely matched and suitable for T. triozae (>20) (Table 2, Figures 20–22).

Table 2. CLIMEX Ecoclimatic Index scores generated by models 1-6 for weather stations in Hawke’s Bay and California. Locality/Region CLIMEX Ecoclimatic Index score for weather station data

Model 1 Model 2 Model 3 Model 4 Model 5 Model 6

Hawke’s Bay (4 weather stations) Napier Taradale Hastings Havelock North Median

54 57 54 0

54

34 37 35 0

34.5

24 32 35 23

28

15 21 23 23

18

58 60 59 53

58.5

36 38 37 31

36.5

California (45 weather stations) Range Median California central valley (10 weather stations) Range Median

0-50 15.2

5-26 20

0-25 7.6

3-10 13

0-50 14.0

9-20 26

0-25 7.6

3-10 13

0-50 19.3

9-26 23.5

0-25 10.0

3-18 11.5

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4 Summary As tomato potato psyllid has a similar range in North and Central America to T. triozae and has established successfully in New Zealand, it is probable that T. triozae can also establish in New Zealand. CLIMEX EI scores suggest that areas of the east coast of the North and South Island are suitable for T. triozae. Caution is needed when interpreting the mapped outputs of the CLIMEX models. Maps of suitable habitat in New Zealand for T. triozae depend on model assumptions about the moisture and temperature preferences of T. triozae. Uncertainty about T. triozae responses to abiotic variables and even the extent to which abiotic variables determine the range of T. triozae in North and central America means that maps produced in this report are necessarily speculative.

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Figure 1. Known distribution of the tomato potato psyllid B. cockerelli (red filled circles), threatened endemic psyllids (light to dark green triangles) and other native Triozid psyllids in New Zealand (pale triangles).

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Figure 2. CLIMEX Ecoclimatic scores generated by model 1 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 1 simulates a warm-dry temperature preference, with cold and wetness limiting distribution.

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Figure 3. CLIMEX Ecoclimatic scores generated by model 2 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 2 simulates a hot-dry preference, with cold and wetness limiting distribution.

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Figure 4. CLIMEX Ecoclimatic scores generated by model 3 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 3 simulates a warm-dry preference with wet limiting distribution.

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Figure 5. CLIMEX Ecoclimatic scores generated by model 4 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 4 simulates a hot-dry preference, with wet limiting distribution.

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Figure 6. CLIMEX Ecoclimatic scores generated by model 5 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 5 simulates a warm-dry preference, with tolerance of wet conditions.

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Figure 7. CLIMEX Ecoclimatic scores generated by model 6 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in North America. Model 6 simulates a hot-dry preference, with tolerance of wet conditions.

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Figure 8. Difference in CLIMEX Ecoclimatic scores generated by model 1 and model 3 for population persistence of the eulophid Tamarixia triozae in North America. Model 1 simulates a warm-dry temperature preference, with cold and wetness limiting distribution. Model 3 simulates a warm-dry preference with wet limiting distribution.

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Figure 9. Difference in CLIMEX Ecoclimatic scores generated by model 3 and model 4 for population persistence of the eulophid Tamarixia triozae in North America. Model 3 simulates a warm-dry preference with wet limiting distribution. Model 4 simulates a hot-dry preference, with wet limiting distribution.

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Figure 10. Difference in CLIMEX Ecoclimatic scores generated by model 4 and model 6 for population persistence of the eulophid Tamarixia triozae in North America. Model 4 simulates a hot-dry preference, with wet limiting distribution. Model 6 simulates a hot-dry preference, with tolerance of wet conditions.

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Figure 11. CLIMEX Ecoclimatic scores generated by model 1 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 1 simulates a warm-dry temperature preference, with cold and wetness limiting distribution. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 12. CLIMEX Ecoclimatic scores generated by model 2 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 2 simulates a hot-dry preference, with cold and wetness limiting distribution. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 13. CLIMEX Ecoclimatic scores generated by model 3 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 3 simulates a warm-dry preference with wet limiting distribution. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 14. CLIMEX Ecoclimatic scores generated by model 4 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 4 simulates a hot-dry preference, with wet limiting distribution. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 15. CLIMEX Ecoclimatic scores generated by model 5 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 5 simulates a warm-dry preference, with tolerance of wet conditions. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 16. CLIMEX Ecoclimatic scores generated by model 6 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 6 simulates a hot-dry preference, with tolerance of wet conditions. Simulation was based on CX_CRU_61-90_V2, an interpolated climate dataset with a spatial resolution of 0.5o (55.6 km at the equator) based on the University of East Anglia Climatic Research Unit CL 2.1 dataset (Mitchell et al. 2004, Stephens et al. 2007). Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 17. CLIMEX Ecoclimatic scores generated by model 2 (based on CLIMEX metdata.mm) indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 2 simulates a hot-dry preference, with cold and wetness limiting distribution. Simulation was based on the CLIMEX climate database metdata.mm, modified by addition of 285 sites for New Zealand. Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 18. CLIMEX Ecoclimatic scores generated by model 4 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 4 simulates a hot-dry preference, with wet limiting distribution. Simulation was based on the CLIMEX climate database metdata.mm, modified by addition of 285 sites for New Zealand. Red circles are localities where tomato potato psyllid has been trapped by PFR staff.

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Figure 19. CLIMEX Ecoclimatic scores generated by model 6 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in New Zealand. Model 6 simulates a hot-dry preference, with tolerance of wet conditions. Red circles are localities where tomato potato psyllid has been trapped by PFR staff. Simulation was based on the CLIMEX climate database metdata.mm, modified by addition of 285 sites for New Zealand.

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Figure 20. CLIMEX Ecoclimatic scores generated by model 2 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in California. Model 2 simulates a hot-dry preference, with cold and wetness limiting distribution.

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Figure 21. CLIMEX Ecoclimatic scores generated by model 4 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in California. Model 4 simulates a hot-dry preference, with wet limiting distribution.

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Figure 22. CLIMEX Ecoclimatic scores generated by model 6 indicating range of unsuitable (0) to optimal localities (>30) for population persistence of the eulophid Tamarixia triozae in California. Model 6 simulates a hot-dry preference, with tolerance of wet conditions.

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5 Acknowledgements We thank Jessica Dohmen-Vereijssen and Nina Jorgensen for supplying the tomato potato psyllid trapping locations in New Zealand. We also thank Esteban Rodriguez Leyva for collection localities within Mexico and copies of relevant papers.

6 References

Gardner-Gee, R 2012. Risks to non-target species from the potential biological control agent Tamarixia triozae, proposed for use against Bactericera cockerelli in New Zealand: A summary of host-range testing. A report prepared for Horticulture New Zealand Ref: SFF09-143 Sustainable TPP management. Plant & Food SPTS No. 7296.

Gomez-Torres ML, Nava DE, Parra JRP 2012. Life table of Tamarixia radiata (Hymenoptera: Eulophidae) on Diaphorina citri (Hemiptera: Psyllidae) at different temperatures. Journal of Economic Entomology 105(2):338–343.

Hamerski MR, Hall RW, Keeney GD 1990. Laboratory biology and rearing of Tetrastichus brevistigma (Hymenoptera: Eulophidae), a larval-pupal parasitoid of the elm leaf beetle (Coleoptera: Chrysomelidae). Journal of Economic Entomology 83(4): 2196–2199.

Ikemoto T 2003. Possible existence of a common temperature and a common duration of development among members of a taxonomic group of arthropods that underwent speciational adaptation to temperature. Applied Entomology and Zoology 38(4): 487–492.

Johnson TE 1971. The effectiveness of Tetrastichus triozae Burks (Hymenoptera: Eulophidae) as a biological control agent of Paratrioza cockerelli (Sulc) (Homoptera: Psyllidae) in north central Colorado. M.S. Thesis, Colorado State University, Fort Collins, Co.

Mitchell TD, Carter TR, Jones PD, Hulme M, New M 2004. A comprehensive set of climate scenarios for Europe and the globe: the observed record (1900-2000) and 16 scenarios (2000-2100). University of East Anglia, 30 pp.

Nechols JR, Tauber MJ, Helgesen RG 1980. Environmental control of diapause and postdiapause development in Tetrastichus julis (Hymenoptera: Eulophidae), a parasite of the cereal leaf beetle, Oulema melanopus (Coleoptera: Chrysomelidae). The Canadian Entomologist 112(12): 1277–1284.

Patil NG, Baker, PS, Pollard GV 1993. Life histories of Psyllaephagus yaseeni (Hym., Encyrtidae) and Tamarixia leucaenae (Hym., Eulophidae),parasitoids of the leucaena psyllid Heteropsylla cubana. Entomophaga 38(4): 565–577.

Pletch DJ 1947. The potato psyllid Paratrioza cockerelli (Sulc): Its biology and control. Bulletin of Montana State College Agricultural Experiment Station. 446: 1–95.

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Rojas PR 2010. Biología de Tamarixia triozae (Burks) (Hymenoptera: Eulophidae) parasitoide de Bactericera cockerelli (Sulc) (Hemiptera: Triozidae). Tesis de Maestría en Ciencias, Colegio de Postgraduados, Texcoco, Edo. de México. 48 p.

Smith AB 2012. The relative influence of temperature, moisture and their interaction on range limits of mammals over the past century. Global Ecology and Biogeography. Article first published online: 19 JUL 2012 | DOI: 10.1111/j.1466-8238.2012.00785.x

Stephens AEA, Kriticos DJ, Leriche A 2007. The current and future potential geographic distribution of the Oriental fruit fly, Bactrocera dorsalis (Diptera: Tephritidae). Bulletin of Entomological Research 97: 369–378.

Sutherst RW 2003. Prediction of species geographical ranges. Journal of Biogeography 30: 805–816.

Sutherst RW, Maywald GF 1991. Climate modelling and pest establishment. Plant Protection Quarterly 6: 3-7.

Tran LT, Worner SP, Hale RJ, Teulon DAJ 2012. Estimating development rate and thermal requirements of Bactericera cockerelli (Hemiptera: Triozidae) reared on potato and tomato by using linear and nonlinear models. Environmental Entomology 41(5): 1190–1198.

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Appendix Table A1. Collection sites for Tamarixia triozae in North and Central America. Location State/Region Country Reference Salvatierra Oaxaca Valles Centrales Jacona de Plancarte Montecillo Texcoco Salamanca, Irapuoto Guanajuato Saltillo Ramos Arispe Culiacán Guasave Marinette 5 miles NW Marinette Phoenix Garden Grove Oxnard, Ventura County Oxnard, Ventura County Orange and Riverside Counties Wildwood Canyon, Yucaipa, San Bernardino County Beaumont, Riverside County San Timoteo Canyon, Riverside County Mill Creek Canyon, San Bernardino County Big Pines Camp, Los Angeles County Wrightwood, San Bernardino County Live Oak Canyon, San Bernardino County Colton, San Bernardino County Irvine, Orange County Weld and Morgan counties Fort Collins Hollister Hobbs Butte Lawrence Billings Billings Billings Bozeman Bozeman Scottsbluff Mesilla Valley Weslaco Lower Rio Grande Valley Spanaway

Guanajuato Oaxaca Oaxaca Michoacan Estado de Mexico Estado de Mexico Guanojuato Guanojuato Guanojuato Coahuila Coahuila Sinaloa Sinaloa Arizona Arizona Arizona California California California California California California California California California California California California California California California Colorado Idaho Idaho Kansas Montana Montana Montana Montana Montana Nebraska New Mexico Texas Texas Washington

Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico Mexico USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA

Rojas 2010 Bravo & Lopez 2007 E. Rodriguez Leyva pers. comm. Lomeli Flores & Bueno Partrida 2002 Rojas 2010 E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm. E. Rodriguez Leyva pers. comm.. E. Rodriguez Leyva pers. comm.. Burks 1943 Burks 1943, GBIF database (type locality) Burks 1943 Burks 1943 Butler & Trumble 2011 Jimenez et al. unpub. report Trumble et al. 2011 Jensen 1957 Jensen 1957 Jensen 1957 Jensen 1957 Jensen 1957 Jensen 1957 Jensen 1957 Jensen 1957 Butler & Trumble 2011 Johnson 1971 Burks 1943 Burks 1943 Burks 1943 Burks 1943 Burks 1943 Burks 1943 GBIF database Burks 1943 GBIF database Burks 1943 Burks 1943 De Leon & Setamou 2010 Yang et al. 2010 Burks 1943

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Bravo ME, Lopez LP. 2007. Principales plagas del chile de agua en los Valles Centrales de Oaxaca. Agroproduce 7:12–15.

Burks BD 1943. The North American parasitic wasps of the genus Tetrastichus. Proceedings of the United States National Museum 93:505–608.

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Butler CD, Trumble JT 2011. New records of hyperparasitism of Tamarixia triozae (Burks) (Hymenoptera: Eulophidae) by Encarsia spp. (Hymenoptera: Aphelinidae) in California. The Pan-Pacific Entomologist. 87(2):130–133.

De León JH, Sétamou M 2010. Molecular evidence suggests that populations of the Asian citrus psyllid parasitoid Tamarixia radiata (Hymenoptera: Eulophidae) from Texas, Florida and Mexico represent a single species. Annals of the Entomological Society of America 103:100–110.

Jensen DD 1957. Parasites of the Psyllidae. Hilgardia 27(2): 71–99.

Johnson TE 1971. The effectiveness of Tetrastichus triozae Burks (Hymenoptera: Eulophidae) as a biological control agent of Paratrioza cockerelli (Sulc) (Homoptera: Psyllidae) in north central Colorado. M.S. Thesis, Colorado State University, Fort Collins, Co.

Lomeli-Flores JR, Bueno Partida R 2002, New record of Tamarixia triozae (Burks), parasitoid of the tomatoe [sic] psilid [sic] Paratrioza cockerelli (Sulc) (Homoptera: Psyllidae) in Mexico. Folia Entomológica Mexicana 41(3):375–376.

Rojas PR 2010. Biología de Tamarixia triozae (Burks) (Hymenoptera: Eulophidae) parasitoide de Bactericera cockerelli (Sulc) (Hemiptera: Triozidae). Tesis de Maestría en Ciencias, Colegio de Postgraduados, Texcoco, Edo. de México. 48 p.

Trumble JT, Butler C, Novy R, Miller C, Kund G, Diaz-Montano J, Carson W 2011. New Approaches for Potato Psyllid Management. Presentation at SCRI Zebra Chip Annual Reporting Session, November 2011.

Yang X; Yong‐Mei Zhang L, Tong‐Xian L 2010. Field and laboratory comparison of life table of potato psyllid (Bactericera cockerelli) and mortality factors analysis on potato and tomato in the Lower Rio Grande Valley of Texas. In: Abstracts of Posters presented during the 64th Annual

Meeting of the Subtropical Plant Science Society (formerly ‐ Rio Grande Valley Horticultural

Society), January 25, 2010; Texas A&M University‐Kingsville, Citrus Center, 312 N.

International Blvd, Weslaco, TX 78596‐9027.