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1 Supplementary Materials for: From Trickle to Flood: The Large-scale, Cryptic Invasion of California by Tropical Fruit Flies Nikos T. Papadopoulos, Richard E. Plant, and James R. Carey * * To whom correspondence should be addressed. jrcarey@ucdavis.edu (JRC) This file includes: Figs. S1-S21 Tables S1-S7 References (69-81)

2 2 FIGURES Fig. S1. Fruit flies detected in California. (top) Worldwide origins of the 17 species of tropical tephritids detected in California including Africa (C. capitata; D. bivitatus), Australia (B. tryoni), Asia (B. albistrigata B. correcta B. cucurbitae B. dorsalis B. oleae B. latifrons B. scutellata B. zonata ), Mexico (A. ludens), and South America (A. obliqua, A. striata, A. suspensa). (bottom) Images of all species excluding B. oleae (olive fly).

3 3 Mexican Fruit Fly (A. ludens) Fig. S2. Detections of the Mexican fruit fly (A. ludens) by period on statewide (upper left), regional (A, B, C), and local (A1, B1, C1) scales. Boxes in A, B, and C designate areas of details in A1, B1, and C1, respectively. Stars indicate locations of initial finds at the state, regional or local scales. The Mexfly was the first tephritid detected in the state having been first discovered in San Ysidro (San Diego County) in There were 4 subsequent outbreaks of this species in San Ysidro over the next 17 years. In early- to mid-1970s the Mexfly began to be recovered in other parts of San Diego County as well as in Los Angeles County. By the mid 1980s it was detected in several cities in Los Angeles, Riverside and Orange Counties, by the mid-1990s they were outbreaks in the Bay Area, and in 2009 a single adult was trapped as far north as Sacramento. In the 57 year period between the first Mexfly capture and 2012 there have been a total of 889 adults and 295 pre-adults recovered on approximately 113 different properties in 77 municipalities in 12 counties. This species has been detected repeatedly a total of 172 times in 25 different cities including 1, 3, 11, and 10 cities with recaptures in 19, 10-12, 3-9, and 2 different years, respectively. Repeat detections in all three of the greater metropolitan areas as well as the Central Coast, Sacramento and San Joaquin Valleys. The Mexican fruit fly is found in Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico and Nicaragua and has been reported from around 50 different host species.

4 4 Melon Fly (B. cucurbitae) Fig. S3. Detections of the melon fly (B. cucurbitae) on statewide, regional, and local scales (see Fig. S2 legend). This species has been captured in 14 different years within the state starting with first detection in the southern California city of Westwood in This was followed by 12 outbreaks 29 to 50 years later including detections ranging from 15 to 50 km of this original find. These subsequent outbreaks were also near each other including the 1999 and 2000 detections in El Monte and Rosemead within 12 to 14 km of the 1985 detections in Pasadena and the 1986 detections in Los Angeles within 7, 14, 5, and 14 km of the detections in Florence (in 1997), Los Angeles (in 1999), Los Angeles (in 2004), and Montebello (in 2006), respectively. In 1985 the melon fly was detected in San Diego and in 2010 it was first detected in the lower Central Valley city of Bakersfield the nearby town of Arvin. Repeat detections have occurred in all three of the greater metropolitan areas as well as the Central Coast, Sacramento and San Joaquin Valleys. There have been multiple recurrent finds in close proximity to original detections within state or region with both northerly and easterly spread. The melon fly is distributed over most of India, which is considered its native home, and throughout most of southeastern Asia. It was introduced into the Hawaiian Islands from Japan about 1895, and by 1897, when it was first observed, it was already a serious pest.

5 5 Oriental Fruit Fly (B. dorsalis) Fig. S4. Detections of the oriental fruit fly (B. dorsalis) on statewide, regional, and local scales (see Fig. S2 legend). First detected in 1960 in Anaheim and Carpenteria in Orange and Santa Barbara Counties, respectively. Whereas detections averaged once every other year during the first 20 years after this species was first detected ( ), outbreaks have occurred every year in the subsequent 33-year period ( ). In the half century since first capture there have been a total of 2,266 adults and 1,705 pre-adults recovered on approximately 1,500 different properties in 245 municipalities in 17 counties. This species has been detected multiple times (n= 369) in 100 different cities including 3 cities with recaptures in 15 to 19 different years, 11 cities with recaptures in 7 to 11 different years, 42 cities with recaptures in 3 to 5 different years, and 44 cities with recaptures in 2 different years. Repeat detections have occurred in all three of the greater metropolitan areas as well in Central Coast regions and in the Central Valley. There were 129 emergency eradication programs directed against B. dorsalis in California between 1982 and This species accounted for over 50% of all emergency eradication programs against fruit flies and over 40% of all eradication programs of any type. Multiple recurrent finds in close proximity to the location of original detections within state or region with spread both north and east. The oriental fruit fly is reported to attack over 230 crops. It is is widespread through much of the mainland of southern Asia and on neighboring islands.

6 6 Guava Fruit Fly (A. striata) Fig. S5. Detections of the guava fruit fly (A. striata) on statewide, regional, and local scales (see Fig. S2 legend). Total of 11 detections, the first of which was in 1963 in San Ysidro. This detection was within 14 km of one 20 years later in San Diego. A 1986 detection in Garden Grove was within 30 to 65 km of outbreaks in Westwood in 1987, Elysian Park in 1989, South Gate in 1998 and Los Angeles in The species has been detected in the two locations in southern California as well as in Fresno in the Central Valley in 1991 and in San Leandro in the Bay Area in This species is found in Mexico (north to southern Sinaloa, Aguascalientes and northern Veracruz) and south to Peru, Bolivia and Brazil. In the West Indies, it is found in Trinidad.

7 7 West Indian Fruit Fly (A. obliqua) Fig. S6. Detections of the West Indian fruit fly (A. obliqua) on statewide, regional, and local scales (see Fig. S2 legend). Although only 8 adult A. obliqua have been captured in the state starting in 1967 with each appearing in different cities in 6 separate years, the reappearances of this species in the state as well as their relatively close proximity suggest that a resident population may exist in the state. There are several specific patterns that support this hypothesis: (1) the initial discovery in 1967 in the Los Angeles district of Wilmington was followed 22 years later in 1989 with 2 appearances in the same month and within around 30 km of the original find and of each other one in Terrance and the other in Anaheim; (2) A third fly was discovered this same year in Chula Vista, San Diego County. (3) Then 3 years later (1992) another adult was discovered only 16 km away in San Diego. (4) This find was followed 11 and 12 years later by discoveries in Corona and Irvine in 2004 and 2005, respectively. These finds were from 40 to 50 km from the detections in 1967 and 1989 and within 22 km of the find 16 years earlier in Anaheim. The distribution of this species includes Mexico south to Argentina, Greater & Lesser Antilles Dominica and St. Lucia. It was briefly established in southern Florida (USA) in the 1930s, but is no longer present. A. obliqua has been recorded from many hosts thought the favored are the mombins, jobos, or hog plums of the genus Spondias, followed by mango, roseapple and guava.

8 8 Mediterranean Fruit Fly (C. capitata) Fig. S7. Detections of the Mediterranean fruit fly (C. capitata) on statewide, regional, and local scales (see Fig. S2 legend). This fly was first detected in 1975 in Santa Monica and several surrounding cities in Los Angeles County. This was followed by recurring outbreaks throughout the 1980s in both southern and northern California. The annual rates of medfly detections increased from 3 out of every 5 years between 1975 and 1989 to 3 out of every 4 years between 1990 and In the 36-year period between first (1975) and last (2012) medfly detections a total of 2,068 adults and 3,884 pre-adults (mostly larvae) were recovered on approximately 1,400 different properties in 170 California municipalities in 21 counties. This species has been detected repeatedly a total of 250 times in 47 different cities including 1 city with medfly recaptures in 11 different years, 2 cities with recaptures in 5 to 6 different years, 12 cities with recaptures in 3 to 4 different years, and 32 cities with recaptures in 2 different years. Repeat detections in all three of the greater metropolitan areas have occurred as well as the Central Coast, Sacramento and San Joaquin Valleys. Multiple recurrent finds in close proximity to original detections within state or region with both northerly and easterly spread since the initial detections in The maps validate the predictions by Carey [68] concerning the pattern and sequence of the invasion in the state including medfly appearances in San Diego followed by outbreaks in the main growing areas of the state including the Central and Imperial Valleys. The Mediterranean fruit fly is native to Africa, but has spread to other parts of the world including southern Europe, Australia, the New World tropics, and Hawaii.

9 9 Caribbean Fruit Fly (B. suspensa) Fig. S8. Detections of the Caribbean fruit fly (B. suspensa) on statewide, regional, and local scales (see Fig. S2 legend). A total of 6 adult Caribbean fruit flies have been captured in California including 3 in 1984 in San Diego, 1 in 1990 in Los Angeles and 1 in each of the years 1996 and 2004 in San Diego and Diamond Bar, respectively. The 1996 find in San Diego was within 5 km of the location of the detection in this city 12 years earlier in Indigenous to the West Indies with a geographic distribution similar to A. oblique and A. striata, this species has a host range of around 100 and is a major citrus pest in Florida where it is established.

10 10 Peach Fruit Fly (B. zonata) Fig. S9. Detections of the peach fruit fly (B. zonata) on statewide, regional, and local scales (see Fig. S2 legend). A native of India and Southeast Asia, the peach fruit fly is has spread to the Middle East and northern Africa. Its host range includes primarily fruits [69]. The peach fruit fly was first detected in 1984 in El Segundo in Los Angeles County. A total of 61 adult flies have been captured in 23 cities located in 9 different counties. Outbreaks have occurred in 12 of the 27 years since the first detection with only two cities experiencing repeat outbreaks. There have been a total of 25 different repeat detections. The 1987 Westchester, 1990 Hollywood and 2009 Anaheim detections were within 5, 20 and 1 km of the 1984 find in El Segundo, respectively. The 1993 detections in Anaheim and Fullerton were within 9 to 15 km of the 2009 detection in Anaheim. The 1998 detection in Pinole in northern California was within 18 km of the detection in Oakland in The 2001 finds in Santa Clara and San Jose were within 9 to 11 km of the 2007 find in San Jose. The 2006 detection in Antelope was within 26 km of the 2010 find in Sacramento. Main areas include Los Angeles Basin (1984), the Inland Empire (2001), Fresno (2006) in the Central Valley, the Bay Area (2001) and Sacramento (2006). Repeat detections in all three of the greater metropolitan areas as well as the Central Coast, Sacramento and San Joaquin Valleys. B. zonata is one of the three most destructive flies in India, causing losses in peach, apricot, guava and figs. Recently it has increased its permanent range to Egypt, Saudi Arabia, Oman, Pakistan, India, Sri Lanka, Mauritius, Reunion Island, Bangladesh, Nepal, Myanmar, Thailand, Laos, Vietnam, and possibly Indonesia (Sumatra).

11 11 Guava Fruit Fly (B. correcta) Fig. S10. Detections of the guava fruit fly (B. correcta) on statewide, regional, and local scales (see Fig. S2 legend). First detected in 1986 in and around Westminster (Orange County), this species has now been found in 59 different cities five primary areas including greater metropolitan areas of Los Angeles and the Bay Area, San Diego, Sacramento, and the Central Valley cities of Modesto and Fresno. It has been detected in the same city or region 103 different times. Repeat detections in all three of the greater metropolitan areas as well as the Central Coast, Sacramento and San Joaquin Valleys. Multiple recurrent finds have occurred in close proximity to original detections within state or region with both northerly and easterly spread since the initial detections in The guava fruit fly occurs in India, Pakistan, Nepal, Sri Lanka, and Thailand and attacks a wide range of fruit host plants.

12 12 Stripped Fruit Fly (B. scutellata) Fig. S11. Detections of the stripped fruit fly fly (B. scutellata) on statewide, regional, and local scales (see Fig. S2 legend). First captured in 1987 in Los Angeles and then again in 19889, 1992, 1993, 1997 and Then in 2009 a major outbreak occurred in and around San Pedro within 5 km radius. Then 3 flies captured the following year (2010). Interestingly and perhaps importantly, unlike virtually all other tephritids which are typically captured in late summer and fall, 11-of-16 A. scutellata detections were in the spring from March through May. Repeat detections in all three of the greater metropolitan areas as well as the Central Coast, Sacramento and San Joaquin Valleys. Multiple recurrent finds in close proximity to original detections have occurred within state or region with both northerly and easterly spread since the initial detections in 1956.

13 13 White striped fruit fly (B. albistrigata) Fig. S12. Detections of the white striped fruit fly (B. albistrigata) on statewide, regional, and local scales (see Fig. S2 legend). A single B. albistrigata adult was captured in one city in 2008 followed by 8 adults in 3 cities in This species occurs on Christmas Island (a territory of Australia), the Andaman and Nicobar Islands (India), Indonesia (Java, Lombok, Sulawesi, Sumatra), peninsular Malaysia, southern Thailand, and probably northern and western Irian Jaya (Indonesia). Its main hosts are Sygygium spp., (e.g. brush cherries from Australia).

14 14 Olive fly (B. oleae) Fig. S13. Detections of the olive fly (B. oleae) on statewide, regional, and local scales (see Fig. S2 legend). First detected in the Los Angeles area in 1998, within 5 years of the first observation this monophagous tephritid was reported from all olive cultivation areas of the state [70]. A native of the Mediterranean Basin it is found throughout this region, in South Africa and now throughout California.

15 15 Anastreapha sp., Bactrocera spp, and Dacus sp, Fig. S14. Detections of: (1) B. facialis (no common name), a polyphagous pest from Tonga with a host list of over 70 species; (2) the solinacious fruit fly, B. latifrons, a pest of solinacious hosts, was discovered in Hawaii in 1983; (3) a highly-polyphagous pest with a host range similar to the oriental fruit fly, the Queensland fruit fly, B. tryoni, is found in Oceania including Australia, French Polynesia, New Caledonia, and Pacific Islands; (4) the African pumpkin fly, D. bivitattus, is found throughout much of Africa attacking mostly cucurbit hosts; and (5) the sapote fruit fly, A. serpentina, is found in Mexico and Central and South America. It attacks a variety of fruits and vegetables although its primary host is sapote fruit. The invasion status of these species is unclear because of the small numbers of captures and/or because they were captured only recently.

16 Fig. S15. Examples of flies captured years apart in nearby locations. Top maps: B. scutellata detections in at locations in Los Angeles 8 years apart within 4 km, and in San Pedro 1 year apart within 4.3 km. Bottom map: B. zonata detections in Garden Grove 6 years apart within 9 km. 16

17 Fig. S16. Examples of flies captured years apart but in close proximity. Top maps: B. dorsalis detections in and around Upland. The four radii from inner to outer are 1, 2, 4, and 8 km in diameter. Note that most of the captures from 1993 through 2008 were within 1 to 4 km of captures in 1987 and Bottom map: B. dorsalis captures 45 years apart within around 10 blocks of each other. 17

18 Fig. S17. Examples of detections years apart but within 2 to 4 km. Top: B. correcta captures in Westminster 11 years apart that were within 2 km; Bottom: B. correcta captures in Garden Grove in 2005 through 2009 that fell within circular regions 4 km in diameter where this same species was detected in 1986 through

19 Fig. S18. Examples of medflies captured years apart within a few hundred meters of earlier locations. Top maps: Relationship of medfly detections in San Jose from 1992 through 2007 relative to captures in this city in during the large outbreak in Santa Clara County. Note in the exploded area of detail that finds were virtually superimposed in the latter period relative to the earlier one and that new medflies were found to the east and south of the original finds. Bottom maps: Same relationship as above except in Mountain View. Medfly finds in 1989 were within 250 to 500 meters of detections during the large Bay Area outbreak in

20 Fig. S19. Seasonality of tephritid (all species) captures in California from 1954 through

21 Fig. S20. Latitudinal (north) distributions of tephritid captures (all species) in California by year. References cities by latitude include San Diego (32.715), Los Angeles (34.052), Fresno (36.748), San Jose (37.304) and Redding (40.587). Note that tephritid captures were restricted to southern California for nearly the first 30 years after the first detection. However, in 1980 with discovery of the medfly in the Bay Area they began appearing in northern California though at the same time reappearing in the south state. The majority of captures in 2000 that lie between 35.0 and 37.0 north latitude represent olive fly detections throughout the Central Valley though several other species also began appearing in this region including B. cucurbitae, B. zonata, and B. dorsalis. 21

22 Fig. S21. Data regarding mitotype of medflies captured in California from 2005 to From the four mitotypes recorded (AAAA, AAAB, AABB, BBBB) one (AAAB) was detected in 78,1% of the individuals tested (n = 114). Earlier studies on the genetic affinity of medfly populations from California and possible areas of origin suggest Guatemala as the possible source of population detected in Southern California, while Hawaii was considered very unlikely source [71-72]. Additional genetic analyses on medfly specimens from the Los Angeles area suggest the existence of an endemic population derived from Guatemala [73]. Results of another study [74] fail to reject the hypothesis of the existence of a resident medfly population in California, and they suggest the introduction of multiple populations from various sources that perhaps overlaid on very localized persistent populations that can occasionally reach detectable levels. Note that the AAAB genotype was found throughout southern California from 2005 through 2010 and also appeared in northern California in 2007 and

23 Table S1A. Regions and sub-regions of the world outside of the U.S. in which the 17 tropical tephritid species captured in California are either endemic (*) or introduced and established (*). Indication of presence in a sub-region does not imply that the species is present in all countries within it. Species designated by first three letters of the species scientific names. Source for world sub-region classification Source for tephritid distributions: [75-78] World Region/Subregion Anastrepha Bactrocera Ceratitis; Dacus lud obl ser str sus alb cor cuc dor fac lat ole scu try zon cap biv Africa Eastern Africa 1 * * * * * 5 Middle Africa 2 * * * * * 5 Northern Africa 3 * * * * 3 Western Africa 4 * * * * * * 5 Southern Africa 5 * * * * * 4 Asia Eastern Asia 6 * * 4 South-Eastern Asia 7 * * * * * * * 7 Western Asia 8 * * * 3 Southern Asia 9 * * * * * * 6 Latin America Caribbean 10 * * * * * 5 Central America 11 * * * * * 5 South America 12 * * * * 3 Oceania Australia 13 * * * 3 Melanesia 14 * * * 3 Micronesia 15 * * * 3 Polynesia 16 * * * * * * 6 Europe Western Europe 17 * * 2 Southern Europe 18 * * 2 1 e.g. Kenya; Tanzania; 2 e.g. Angola; Congo; 3 e.g. Morocco; Egypt; 4 e.g. Sierra Leone; Benin; 5 e.g. South Africa; Botswana; 6 e.g. China; Japan; 7 e.g. Thailand; Indonesia; Philippines; 8 e.g. Turkey; Israel; 9 e.g. India; Afghanistan; 10 e.g. Bahamas; Puerto Rico; 11 e.g. Costa Rica; Mexico; 12 e.g. Colombia; Brazil; 13 e.g. Australia; New Zealand; 14 e.g. Fiji; Papua New Guinea; 15 e.g. Guam; Marshall Islands; 16 e.g. Tonga; French Polynesia; 17 e.g. France; Austria; 18 e.g. Spain; Greece TOT

24 24 Table S1B. Confirmed or likely host species for each of the 17 tropical tephritid species detected in California. Sources from references contain in the Literature Cited of Aluja and Mangan (2008). Hosts Anastrepha Bactrocera Ceratitis; Dacus lud obl ser str sus alb cor cuc dor fac lat ole scu try zon cap biv Fruits (Genus) Almonds (Prunus) * 1 Apples (Malus) * * * * * * 6 Avocado (Persea) * * * * * * 6 Berries (Vaccinium) * * 2 Cherries (Prunus) * 1 Citrus (Citrus) * * * * * * * * * * * 11 Figs (Ficus) * * * * * 5 Grapes (Vitis) * 1 Olives (Olea) * * 2 Peaches (Prunus) * * * * * * * 7 Pears (Pyrus) * * * * * * * 7 Plums (Prunus) * * * * * 5 Vegetables (Genus) Cucumber (Cucumis) * * * 3 Eggplant (Solanum) * * * * 4 Melons (Cucumis) * * * * * 5 Peppers (Capsicum) * * * * * * 6 Squash (Cucurbita) * * * * * 5 Tomatoes (Solanum) * * * * * * * * 8 Tropical (Genus) Mango (Mangifera) * * * * * * * * * * * * * 13 Papaya (Carica) * * * * * * * 7 Guava (Psidium) * * * * * * * * * * * 11 TOT

25 Table S2. Tephritid eradication projects in California by year and species. (SOURCE: California Department of Food and Agriculture). Species Year Medfly Oriental Peach Melon Mexfly Carab Guava Totals

26 Table S3. Interceptions of tephritid fruit flies in airline baggage at eight international airports in the U.S. Although the more northerly cities are not located in fruit fly-friendly areas, data from these cities is included to illustrate the pervasiveness of tephritid interceptions at all airports throughout the U.S. [79] California Fruit Fly-Friendly Regions Fruit Fly-Unfriendly Regions Hawaii Year Los Angeles Dallas Miami Atlanta Boston Chicago New York Honolulu Total Annual

27 27 Table S4. Insect interceptions at ports of entry and border stations over a 17-year period from 1984 to 2000 in states or regions vulnerable to tephritid fruit fly [80]. Note that interceptions in California (i.e. San Diego, Los Angeles, and San Francisco) were 16-18% of the total for cities in each of the subgroups. Cities State Number insects intercepted CITIES NEAR MEXICAN BORDER Laredo TX 17,125 El Paso TX 19,054 Brownsville TX 11,285 Hildalgo TX 5,898 Eagle Pass TX 894 Nogales AZ 8,551 San Diego CA 11,699 74,506 CITIES INTERIOR OR NOT NEAR BORDER Houston TX 21,722 Dallas TX 12,681 Honolulu HI 48,965 Erlanger KY 2,232 New Orleans LA 5,719 San Juan PR 45,965 Miami FL 117,498 Ft Lauderdale FL 10,221 Port Orlando FL 2,162 Atlanta GA 8,437 Savannah GA 2,160 Los Angeles CA 52,666 San Francisco CA 9, ,860

28 28 Table S5. Tephritid fruit fly interceptions from in 2007 through 2009 at Charles de Gaulle International Airport (Paris) by the French National Plant Protection Organization. Unlike tephritid interceptions in the U.S. which can only be identified to family or subfamily because larvae cannot be reared to adults, this interception data identifies species because all intercepted larvae were reared to adults. SOURCE: Balmès Valérie, Laboratoire National de la Protection des Végétaux, Station de Montpellier. CBGP Campus International de Baillarguet CS, YEAR Source/Source Totals Anastrepha spp. Brazil Mexico Peru Domin. Rep TOTAL B. correcta Thailand Vietnam TOTAL B. cucurbitae Maurtius Thailand TOTAL B. dorsalis Cambodia India Pakistan Phillipines Sri Lanka Thailand Vietnam TOTAL B. invadens Burkina Faso Cameroon Ivory Coast Kenya Mali Centrafican Rep Senegal Togo TOTAL

29 29 Table S5 (continued). YEAR Source/Source Totals B. latifrons Thailand Vietnam TOTAL B. zonata Egypt India Pakistan Thailand Vietnam TOTAL C. capitata 0 South Africa Brazil Israel Lebanon Peru TOTAL C. cosyra Burkina Faso Ivory Coast Kenya Mali Senegal TOTAL GRAND TOTALS

30 30 Table S6. Tephritid and total number of interceptions in EU countries and Switzerland in Source: EUROPHYT is a web-based network and database connecting Plant Health Authorities of the EU Member States and Switzerland, the European Food Safety Authority and the Directorate General for Health and Consumers of the European Commission. World Region of Export Country Total Interceptions Tephritid Interceptions Number Percent of Total Africa Burkina Faso Cameroon Congo Cote Divoire Egypt Ghana Guinea Guinea Bissau Kenya Madagascar Mali Mauritius Senegal South Africa Sudan Uganda Asia/Mideast Bangladesh Cambodia China India Laos Malasia Pakistan Philippines Saudi Arabia Sri Lanka Thailand Viet Nam Americas Brazil Colombia Domincan Republic Jamaica Peru United States Total 1,

31 31 Table S7. Establishment status (i.e. long-term, reproducing resident population) ascribed to the 17 tephritid fruit fly species detected in California. Excepting the olive fly which is considered by both state and federal agencies to be established in the state, each species was assigned to a category of establishment probability based on: (i) historical absence or near absence of detections in other fruit fly friendly regions; (ii) outcome of statistical analyses/modeling for a subset of species (i.e. A. ludens, B. dorsalis, C. capitata); and (iii) combination of invasion metrics including year span, total number of years captured, capture frequency, number of individuals caught, and geographic spread based on the number of cities detected. The four categories (CAT) include: I=established; II=establishment virtually certain; III= establishment likely; IV=establishment status unknown. Tephritid Species # Species Common Name Establishment Status First Year CAT Invasion metrics 1 B. oleae Olive fly 1998 I Widespread; large populations state-wide 2 A. ludens Mexican fruit fly 1954 II 3 B. dorsalis Oriental fruit fly 1960 II 58 year span; 41 years captured; 77 cities; 1,200 flies; 113 detections 52 year span, 44 years; 244 cities; > 4,000 flies; > 1,500 detections 4 C. capitata Mediterranean fruit fly 1975 II 37 year span; 26 years and 169 cities; nearly 5,000 flies; 1,400+ detections 5 B. correcta Guava fruit fly 1986 II 6 B. cucurbitae Melon fly 1956 III 7 A. striata Guava fruit fly 1963 III 8 B. zonata Peach fruit fly 1984 III 9 B. scutellata Striped fruit fly 1987 III 26 year span; 23 years and 59 cities; 131 flies 55 year span, 15 years; 17 cities; extremely persistent; 29 detections; 34 flies 48 year span; 11 cities; 10 detections and 11 flies in 9 years 26 year span, 12 years, 57 detections; 23 cities; 61 flies in 57 cities 33 years span; 6 years; 15 detections and 15 flies in 8 cities

32 32 Table S7 (continued) Tephritid Species # Species Common Name Establishment Status First Year CAT Invasion metrics 10 A. obliqua West Indian fruit fly 1967 IV 11 A. suspensa Caribbean fruit fly 1983 IV 42 year span; low number of detections; 7 flies in 8 cities 21 year span; low captures; 6 flies in 3 cities over 4 years 12 B. tryoni Queensland fruit fly 1985 IV 2 flies in 2 cities 13 D. bivitattus African pumpkin fly 1987 IV Single fly, single detection, single city 14 A. serpentina Sapote fruit fly 1989 IV Single fly, single detection, single city 15 B. latifrons Malasian fruit fly 1998 IV Single fly, single detection, single city 16 B. facialis No common name 1998 IV Single fly, single detection, single city 17 B. albistrigata White striped fruit fly 2009 IV Total of 8 flies, in 3 detections in 3 cities

33 33 REFERENCES 69. Carey JR The future of the Mediterranean fruit fly population in California: a predictive framework. Biol. Cons. 78, Qureshi A, Hussain T, Carey JR, Dowell RV Effects of temperature on development of Bactrocera zonata (Saunders) /(Diptera: Tephritidae). Pan-Pacific Entomol. 69, Zygouridis NE, Augustinos AA, Zalom FG, Mathiopoulos KD Analysis of olive fly invasion in California based on microsatellite markers. Heredity 102, Haymer DS, He M, McInnis DO Genetic marker analysis of spatial and temporal relationships among existing populations and new infestations of the Mediterranean fruit fly (Ceratitis capitata). Heredity 79, He M, Haymer DS Genetic relationships of populations and the origins of new infestations fo the Mediterranean fruit fly. Molec. Ecol. 8, Bonizzoni M, Zheng L, Guglielmino CR, Haymer DS, Gasperi G, Homulski LM, Malacrida AR Microsatellite analysis of medfly bioinfestations in California. Molec. Ecol.10, Meixner MD, McPheron BA, Silva JG, Gasparich GE, Sheppard WS The Mediterranean fruit fly in California: evidence for multiple introductions and persistent populations based on microsatellite and mitochondrial DNA variability. Molec. Ecol.11, White IW Taxonomy of the Dacina (Diptera: Tephritidae) of Africa and the Middle East. Afri. Entomol. Memoir 2, Robinson AS, Hooper G Fruit Flies Their Biology, Natural Enemies and Control. (Amsterdam, Elsevier.

34 Bateman MA The ecology of fruit flies. Annu. Rev. Entomol. 17, Fletcher BS The Biology of Dacine Fruit Flies. Annu. Rev. Entomol. 32, Liebhold AM, Work TT, McCullough DG, Cavey JF Airline baggage as a pathway for alien insect species invading the United States. Amer. Entomol. 52, McCullough DG, Work TT, Cavey JF, Liebhold AM, Marshall D Interceptions of nonindigenous plant pests at US ports of entry and border crossings over a 17-year period. Biol. Invas. 8,

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