Report of the

Tomato Genetics Cooperative

Volume 60 December 2010

2

Report

of the

Tomato Genetics Cooperative

Number 60- December 2010 University of Florida

Gulf Coast Research and Education Center

14625 County Road 672

Wimauma, FL 33598 USA

Foreword

The Tomato Genetics Cooperative, initiated in 1951, is a group of researchers who share and interest in tomato genetics, and who have organized informally for the purpose of exchanging information, germplasm, and genetic stocks. The Report of the Tomato Genetics Cooperative is published annually and contains reports of work in progress by members, announcements and updates on linkage maps and materials available. The research reports include work on diverse topics such as new traits or mutants isolated, new cultivars or germplasm developed, interspecific transfer of traits, studies of gene function or control or tissue culture. Relevant work on the Solanaceous species is encouraged as well. Paid memberships currently stand at approximately 94 (includes those paid in 2009 and beyond) from 16 countries. Cover: Design by Dolly Cummings. Bacterial wilt incited by Ralstonia solanacearum is a serious threat to tomato production in many humid tropical production regions. Breeding for resistance has been a challenge due to multiple strains of the bacterium, variable environmental effects on disease expression, largely unknown genetics for resistance, and linkage of resistance to small fruit size. Locating molecular markers tightly linked to resistance genes should be a boon to future breeding efforts and with the tomato genome now sequenced this research may advance much more rapidly than in the past. This years‟ feature article explores the major sources (“roots”) of bacterial wilt resistance and thereby sheds light on the relationships of genotypes that could be used in studies to locate molecular markers. Precision in the exact identification of sources is hampered by a lack of information or by conflicting information. Unfortunately some of the old sources are no longer available. If you read the article and have information that would help us improve the article please contact us. We live in an electronic age and if we get better information we will put it in the article and future versions may be better than the one published in December of 2010. If you have seed of a rare source line send it to us and we will see that it gets in a gene bank. Thanks for your help.

3

Table of Contents

Foreword 2

Announcements 4

Feature Article

Tomato resistance to bacterial wilt caused by

Ralstonia solanaearum E.F. Smith: ancestry and peculiarities

Daunay M.C., Laterrot H., Scott J.W., Hanson P., Wang J.-F 6

Fig 1: Origins Tomato Bacterial Wilt material 20

Table 1: Summing up of the phenotype of some breeding lines 21

Fig 2-12: Pedigree Montage 30

Research Reports

Preliminary Observations on the Effectiveness of Five

Introgressions for Resistance to Begomoviruses in Tomatoes

Luis Mejía, Rudy E. Teni, Brenda E. García, Ana Cristina Fulladolsa,

and Luis Méndez; Sergio Melgar, and Douglas P. Maxwell 41

Preliminary report on association of ‘Candidatus Liberibacter

solanacearum’ with field grown tomatoes in Guatemala

Luis Mejía, Amilcar Sánchez, and Luis Méndez; D. P. Maxwell;

R. L. Gilberston; V.V. Rivera and G.A. Secor 54

Study of epidermal cell size of petals and stamens in tomato

species and hybrids using confocal laser-scanning microscopy

Christopher Lofty, Julian Smith, Pravda Stoeva-Popova 58

Stock Lists 66

Membership List 101

Author Index 107

4

Announcements From the editor:

Help I‟ve fallen behind and I can‟t catch up! The 2010 TGC is late as a result but it is still 2010 so it could be worse- my apologies for the delay. This is our first year of our “electronic only” (see below) format and we are determining how to proceed. No dues were requested from the members in 2010 as most (but not all) costs are associated with printing and mailing the report. On the web we will post only the Table of Contents for a year and will mail a link to an electronic version of Volume 60 to members who paid in 2009. We can also send a printed version to those who want one and will pay to have one sent. Members will receive an email about this option. The cost would be $20US for domestic members and $25US for foreign members. Make checks payable to The University of Florida from a US bank or a bank with a US affiliation. Sorry no credit cards can be used. If you do not have easy access to a bank with a US affiliation we can accept cash in US dollars. For those who only want the electronic version we will ask for dues of $10 per year starting in 2011. Members will receive an email about this in spring 2011 but send in your dues at any time, either for electronic only or for electronic and printed versions as per the prices stated above.

I have not been happy with the key word search of the TGC Reports available on our website: (http://tgc.ifas.ufl.edu/) as it picks up words in areas outside of the reports such as from the Table of Contents and thus is somewhat messy. We have discussed a way to fix this and hope to have it fixed in 2011.

You can see that there are only 3 research reports in this volume. This epitomizes the trend we have been seeing over the last several years as researchers are not sending in reports. Perhaps this year‟s dearth of reports is due in part to the change to an electronic only format. However, I do see a place for the TGC here in the 21st century and plan to keep moving forward. I hope you will help by retaining your membership or becoming a member if you are not presently one and by sending in reports, varietal pedigrees etc.

Last but certainly not least, my heartfelt thanks to Dolly Cummings who keeps TGC business in order around here. Thanks to Dolly and Christine Cooley who work on the website updates.

My contact information: Jay W. Scott, Ph.D. Gulf Coast Research & Education Center 14625 CR 672 Wimauma, FL 33598 USA Phone: 813-633-4135; Fax: 813-634-0001 Email: jwsc@ufl.edu Jay W. Scott Managing Editor

5

Upcoming meetings: February 17-19, 2011, Sol-Conference 2011 Chiangmai, Thailand http://www.sol-symposium2011.com/abstra.aspx March 20-23, 2011 43rd Tomato Breeders Roundtable * El Cid Castilla Beach Resort Hotel, Mazatlan, Sinaloa,, Mexico. http://tgc.ifas.ufl.edu/2011TBR.htm April 11-14 2011 XVIIth EUCARPIA Meeting - Section Vegetables - Working Group Tomato, Málaga, Spain http://www.eucarpiatomato2011.org October 16-20, 2011 SOL & ICuGl Joint Conference 2011 Tsukuba International Congress Center (EPOCHAL), Tsukuba, Japan http://www.sol2011.jp * Recently cancelled-might be held in conjunction with the Tomato Disease Workshop in October at Cornell University, date not yet set nor is it official yet so check for updates on the TGC website at http://tgc.ifas.ufl.edu .

6

FEATURE ARTICLE TGC REPORT VOLUME 60, 2010 Tomato resistance to bacterial wilt caused by Ralstonia solanaearum E.F. Smith: ancestry and peculiarities

Daunay M.C. (1), Laterrot H. (1bis), Scott J.W. (3), Hanson P. (4), Wang J.-F (4).

(1) INRA, UR 1052, Montfavet, France, (1bis) retiree of INRA, UR 1052 (2) CIRAD, Pôle 3P, Saint Pierre, Réunion Island, France (3) University of Florida, Gulf Coast Research & Education Center, Wimauma,

Florida, USA (4) AVRDC-The World Vegetable Center, Tainan, Taiwan

Summary Several national tomato breeding projects began work on developing varieties

resistant to bacterial wilt over 60 years ago and several varieties created in the 1950s, 1960s, 1970s and later on are still found as reference varieties in many recent publications dealing with the genetics of resistance. From the beginning there were many exchanges of resistant material between the breeding programs that are difficult to retrace because published information is scarce. As a consequence the source(s) of resistance of the reference varieties, and the relationships between these varieties are often unclear. This paper provides a synthesis of the relationships between the breeding carried out in Puerto Rico, the USA (North Carolina, Hawaii, Florida), Japan, the Philippines, the French West Indies, and Taiwan, the main sources of resistance that they used, as well as the parentage between the lines they created. The limits of the reliability of our results are explained. The information on the resistance of many bacterial wilt resistant lines to other vascular diseases is also summarized together with some other peculiarities, in order to provide a synthesis useful for breeding bacterial wilt resistant tomatoes and for further genetic studies of the resistance patterns.

Introduction Bacterial wilt is caused by the pathogen formerly known as Pseudomonas

solanacearum, transitorily renamed Burkholderia solanacearum (Yabuuchi et al., 1992) and presently accepted as Ralstonia solanacearum (Yabuuchi et al., 1995; Vaneechoutte et al., 2004). Developing varieties with resistance has challenged tomato breeders for over 60 years for several reasons. Strong interactions are observed between resistance, environmental conditions and strains (e.g. Kelman, 1953; Acosta, 1963; Krausz & Thurston, 1975; Messiaen, 1989; Peter et al., 1993; Prior et al., 1994; Hanson et al., 1996; Jaunet & Wang, 1999; Balatero et al., 2005; Hai et al., 2008). Further, several defaults are often associated to bacterial wilt resistance, such as small fruit size (Acosta et al., 1964; Gilbert & Tanaka, 1965; Scott et al., 2005), bitterness due to high tomatine content (Borchers & Nevin, 1954; Mohanakumaran et al., 1967; Digat & Derieux, 1968; Messiaen, et al., 1978), green gel around the seeds and epidermis cracking (Acosta, 1963; Cordeil & Digat, 1967; Daly, 1976). The difficulty in developing resistance with adaptation to heat and with good horticultural features, in particular fruit firmness and large size, has been reported by many authors (e.g. Kaan et al., 1969;

7

Opena et al., 1989; Celine et al., 2003; Scott, 1997; Scott et al., 2003). It is also very difficult to combine bacterial wilt resistance to resistance to root knot nematode (Messiaen et al., 1978; Prior et al., 1994; Deberdt et al., 1999 a & b).

Kelman (1953) and Acosta (1963) reported the first surveys of the early screening trials carried out during the first half of the 20th century, in the USA and other countries, with hundreds of tomato varieties. The very limited success obtained at that time outlined the difficulty to identify highly resistant material. The breeding for resistance took on new momentum after World War II. Sources of high levels of resistance were identified and used for breeding. The material most widely used nowadays as; controls, key source(s) of resistance for ongoing breeding programs and/or for genetic studies was created in the span of time running from the 1950s and 1970s. Our main objective was to draw a worldwide historical outline of the major breeding inputs of this period, in order to get a global picture of the key original resistance sources used, of the major breeding lines obtained, and of their relationships. Indeed, as information on these topics is scarce, scattered and confusing, it is useful to sum it up for the sake of present and future research on tomato resistance to bacterial wilt. The major programs were carried out (1) in Puerto Rico, (2) in US Universities (North Carolina, Hawaii, Florida), (3) in the Horticulture Research Station1, Japan, (4) in the University of Philippines College of Agriculture, (5) in the French public institutes INRA2 in Guadeloupe and IRAT3 in Guadeloupe and Martinique (French West Indies), and later on (6) in AVRDC4, Taiwan.

1. Primary historical breeding programs, their sources of resistance and germplasm flow between programs

The main accessions used and created in the major research and breeding programs of the USA, Japan, Philippines, French West Indies and Taiwan, as well as their relationships, are outlined in Fig. 1. For the sake of clarity the accessions listed are limited to the major ones, i.e. those which are most frequently mentioned in the literature for their high level of resistance in worldwide trials, or for which the sources of resistance have been published. We also took care to mention enough accession names for the reader to obtain the encoding system used by the different programs. The arrows linking two accession names indicate their parental link. The arrows starting from the frame of a given program indicate the use of material of this program in another program or for creating a given breeding line. The number(s) in brackets indicate the literature reference where the information about the relationships between accessions and programs is provided. For the convenience of the reader, we summed up in Table 1

1 The name and locations of this institute changed along the time: 1921-1950: Horticulture Research

Station (Okitsu, Shizuoka); 1950-1961: Department of Horticulture, National Institute of Agricultural Sciences (Hiratsuka, Kanagawa); 1961-1973: Horticulture Research Station (Hiratsuka, Kanagawa); 1973-1986: Vegetable and Ornamental Crops Research Station (Tsu, Mie); 1986-2001: National Research Institute of Vegetables, Ornamental plants and Tea (NIVOT) (Tsu, Mie); 2001-present: National Institute of Vegetable and Tea Science (NIVTS) (Tsu, Mie). Dr H. Fukuoka, NIVTS, pers. com. 2 Institut National de la Recherche Agronomique.

3 Institut de Recherche en Agronomie Tropicale (now part of CIRAD, Centre de Coopération

Internationale en Recherche Agronomique pour le Développement). 4 The Asian Vegetables Research and Development Centre (now AVRDC-The World Vegetable Center).

8

available information on the phenotype of the main resistant accessions lines displayed in Figure 1 and issued from the main historical breeding programs. Puerto Rico

Information about tomato breeding for bacterial wilt resistance in the University of Puerto Rico is very scarce and dispersed in annual reports of the Puerto Rico University Agricultural Experiment Station. For instance, Cook (1934, 1935), Roque (1935) and Theis (1950) mention partial resistance of some native material and its use in breeding. Warmke & Cruzado (1949) experimented with local selections from hybrids between native and imported tomato varieties, some of which showing some resistance to bacterial wilt and out-yielding the controls. Azzam (1964) reported the existence of resistance in S. pimpinellifolium as well as the development of breeding lines with some degree of resistance but with unacceptable fruit quality.

At the beginning of the 1960s, at Rio Piedras station, a double cross involving [„Platillo‟,( a native variety) X a S. pimpinellifolium, (of unknown identity)] X [a tomato, (of unknown identity) X „Platillo‟] was made by H. Azzam (Digat & Derieux, 1968; Daly, 1976) and its progenies were used by IRAT in the 1960s and 1970s [see below]. The literature also mentions „Beltville 3814 (=T414)‟ which was said to be from Puerto Rico and was used in North Carolina breeding effort [See below].

North Carolina The search and breeding of bacterial wilt resistant tomatoes began long ago in the

USA, at the turn of the 19th and 20th century in the agricultural experiment stations of several States including North Carolina, Florida, Alabama, and Mississipi (Kelman, 1953). After a lapse of years, efforts were resumed at the North Carolina Experiment Station in 1936 (Schmidt, 1936, 1937) and involved many horticultural scientists such as W.S. Barham, F.D. Cochran, M.E. Gardner, W.R. Henderson, J.S. Weaver, and plant pathologists such as D.E. Ellis, S.F. Jenkins, A. Kelman and N.N. Winstead (Henderson & Jenkins, 1972b). Warmke & Cruzado (1949) as well as Walter (1967) mention the existence of a US Southern Tomato Exchange Program (STEP) that was put into operation in 1946 (Yarnell, 1948), and was complemented with the National Screening Program for evaluation of PI (Plant Introduction) accessions of Lycopersicon for disease resistance. These programs are probably at the origin of the complex relationships between the breeding research carried out in North Carolina, Hawaii, Florida and Puerto Rico for bacterial wilt resistance.

The two widely mentioned sources of resistance of North Carolina breeding material are S. lycopersicum var. cerasiforme „PI 129080‟ (=T 702) from Colombia (initially classified as L. pimpinellifolium -Henderson & Jenkins, 1972b) and (ii) S. lycopersicum var. pyriforme „Beltsville No. 3814‟ (=T 414) (Henderson & Jenkins, 1972a & b; Laterrot et al., 1978; Hanson et al., 1998). „Beltsville No. 3814‟ also named „P.I. No. 3814‟ by Kelman (1953) originated in Puerto Rico according to this author and others (Winstead & Kelman, 1952; Henderson & Jenkins, 1972a & b) without further details. Thurston (1976) also said that it is a selection from Puerto Rico. However the name of this line and the fact that it has a PI number both suggest that „Beltsville No. 3814‟ was associated with the USDA, Beltsville (Maryland). Perhaps USDA researchers were collaborating with Puerto Rican researchers. „T414‟ displayed, as other bacterial wilt

9

resistant lines, a specific bitter taste with lasting burning sensation, and it was used by Borchers & Nevin (1954) for setting up a quantitative chemical test of the alkaloids responsible of this taste. Another source of resistance, S. lycopersicum „Mulua‟ from Guatemala, is mentioned by Winstead & Kelman (1952) and by Suzuki et al. (1964) [who refer to Winstead & Kelman (1952) as well as to a personal communication of D.E. Ellis]. According to Winstead & Kelman (1952), „Mulua‟ yielded resistant breeding material after an initial cross with „Rutgers‟. Suzuki et al. (1964, p.99) still referring to Winstead & Kelman (1952), mention also a „T-141‟ from Puerto Rico as another source of resistance used in North Carolina, though these latter authors mention „T414‟ and not „T-141‟. The identity of „T-141‟ is henceforth doubtful, either a mistyping of „T414‟ by Suzuki et al. (1964) or another line not reported in any other source we found.

To sum up, the cherry tomato „PI 129080‟ (=T 702) from Colombia, the pear shaped tomato „Beltsville No. 3814‟ (=T 414) and the tomato „Mulua‟ from Guatemala, have been included in NC breeding programs as genitors of bacterial wilt resistance.

„Venus‟ and „Saturn‟, released in the early 1970s (Henderson & Jenkins, 1972a & b) are the best known commercial varieties with resistance to bacterial wilt issued from the North Carolina State program. Their pedigrees are provided in Fig. 2, and their phylogenic relationship with two other NC lines, „MR4‟ and „NC 72 TR 4-4‟, is provided in Fig.3.

Hawaii D.C. McGuire, J.C. Gilbert and J.C. Acosta (breeders) as well as I.W. Buddenhagen

(pathologist) are among the scientists having worked on tomato resistance to bacterial wilt in the course of the 1950s and 1960s. Breeding for resistance in commercial type tomatoes was confined first (prior to 1955) to crossing root knot nematode resistant Hawaii lines and North Carolina bacterial wilt tolerant lines (Acosta et al., 1963, 1964). Acosta (1963) indicated that several North Carolina lines which had been bred for bacterial wilt resistance, proved to be intermediate in wilt susceptibility under Hawaiian conditions. In 1953 a new source of resistance, S. pimpinellifolium „PI 127805A‟, originating from Peru, was “obtained” [sic]5 and field selected in Hawaii through 9 generations (Acosta et al., 1964). This accession would be at the origin of the line „5808-2‟ (Mohanakumaran et al., 1967). Acosta (1963) writing the name as „HES 5808-2‟, mentions that it is an inbred line of L. pimpinellifolium obtained by D.C. McGuire, but he does not refer to any PI number. The commercial variety „Kewalo‟ (Fig. 4), developed by Gilbert et al. (1974), recombines the resistance to bacterial wilt originating from „PI 127805A‟ with root-knot nematode resistance(gene Mi) and other useful traits from „Anahu‟, a local tomato, and its derivative „Kalohi‟.

„Hawaii 7996‟ (H7996), „Hawaii 7997‟ (H7997) and „Hawaii 7998‟ (H7998) were later bred by J.C. Gilbert in the 1970s (Scott et al., 2005) and it has been reported in the literature (Hanson et al., 1998; Balatero & Hautea, 2001; Scott et al., 2005) that these lines had resistance derived from PI 127805A. However, correspondence dated October 1983 from J. Tanaka (Assistant Horticulturist at Hawaii University) to J. Scott indicates that the 3 mentioned Hawaiian lines and five others („H7975‟, „H7976‟, „H7981‟, „H7982‟, „H7983‟) are sister lines selected from a initially highly variable

5 The word “obtained” in this sentence is quite imprecise, since it might mean “received from someone” or

“obtained by breeding”.

10

accession named „HSBW‟, which acronym might mean „Hot Set Bacterial Wilt‟. This enigmatic accession, delivered to J.C. Gilbert via an unidentified way and planted first in 1973, displayed a high level of bacterial wilt resistance in hot tropical areas. An earlier correspondence, dated March 1978, from J.C. Gilbert to H. Laterrot (INRA) indicates that „Hawaii 7996‟ could be cited as having been selected at the University of Hawaii for bacterial wilt resistance, and as having its origin somewhere in the Philippines. J.C. Gilbert admitted in this letter that no publications had been specifically written for this line, and that he was not fully satisfied with it because of its flavour due to alkaloid residues in the ripe fruit. He recommended to using it as a rootstock or as a parent to be crossed with another parent of good flavour and some bacterial wilt resistance, for making F1s. Laterrot et al. (1978) also mentioned as a personal communication of Gilbert, a Philippine source at the origin of „Hawaii 7996‟. H. Laterrot recorded in his handwritten notes, based on Kaan personal communication, that „Hawaii 7996‟ was a selection made in „Hotset X Philippine tomato‟ and that it had very small fruits and a determinate growth habit. Given the absence of further written details, it is probably impossible to unravel further the pedigrees of „HSBW‟ and „Hawaii 7996‟.

In conclusion, a close look at the dispersed information relative to the breeding programs of Hawaii University indicates that several sources of resistance have been used successively there, first North Carolina material, then „PI 127805A‟ and lastly Philippines material. It is possible (or probable), that J.C. Gilbert recombined these sources along the time in his breeding material, in one way or another. Breeding is an art as much as a science, and the exact pedigree of the most famous bacterial wilt resistant line „Hawaii 7996‟ will probably remain the secret of the breeding genius of the late J.C. Gilbert.

Florida According to Sonoda et al. (1979) the first attempts to search for bacterial wilt

resistance in Florida started over one century ago, but the breeding efforts took a real momentum in the late 1970s. The original sources of resistance used were „Hawaii 7997‟, S. lycopersicum var. cerasiforme „CRA 66‟ and S. lycopersicum „PI 126408‟. The latter is one of the 28 PI accessions determined as resistant to bacterial wilt out of 909 accessions tested (Barham & Ellis, 1951). However most of the material derived in Florida utilized „Hawaii 7997‟. Attempts to pyramid resistance genes in the early 1980‟s were not successful as there were no molecular markers to identify the genes in resistant plants. No lines were developed that had resistance greater than any of the sources, so there was no evidence that improvements were made under Florida conditions. Over the years it was evident that it was difficult to attain large fruit with high resistance levels. In 1995 „Neptune‟ (Fig.5), a line with larger fruit size than „Hawaii 7997‟, was released, but its resistance level was less than that of „Hawaii 7997‟ (Scott et al., 1995a), and when tested in the world wide test (as Fla. 7421) „Neptune‟ also displayed a much narrower spectrum of resistance (Wang et al., 1998). Breeding efforts then focused on taking lines like „Neptune‟ and crossing them back to „Hawaii 7997‟ to attain large fruited lines with high resistance levels. At first, new lines were developed with moderate fruit size and high resistance. The next crosses were with very large fruited susceptible inbreds. From this work the sister lines „Fla. 8109‟ and „Fla. 8109B‟ were developed (Scott et al., 2003) and further crossing provided new inbreds with high

11

resistance and very large fruit (Scott et al., 2009). Hypothetically, „Fla. 8109‟ and the lines developed thereafter contain a resistance gene, missing in „Neptune‟, that was unlinked from a gene preventing large fruit formation, but this has not been elucidated yet.

Japan In Japan, development of bacterial wilt resistance in tomato (and eggplant) started

as early as 1951 and was based on the use of North Carolina lines. „OTB-1‟ and „OTB-2‟ are self pollinated offspring obtained in the 1950s respectively from „NC1953-60N‟ and „NC1953-64N‟ (Suzuki et al., 1964). According to later reports and papers published [in Japanese] by the Ministry of Agriculture and Forestry (H. Fukuoka6, pers. com.) „OTB-2‟ was segregating for several traits including bacterial wilt resistance (but was fixed for Fusarium wilt resistance) and was submitted to further screening for bacterial wilt resistance and further selfing. In 1969 „BF-Okitsu 101‟ was obtained7 from this process. „OTB1‟ and „OTB2‟ were described in 1964 by IRAT (French West Indies) as possessing the traits of the Puerto Rican S. pimpinellifolium, but with markedly bigger fruits and exceptional fruit productivity, together with a good behaviour towards viruses.

Philippines In Philippines, breeding was established as early as 1954 by T.L. York and J.R.

Deanon who evaluated local and foreign accessions with known resistance to bacterial wilt (Deanon, 1988). The exact origin of the resistance source(s) used in the Philippines breeding scheme is not found in the literature and hence remains confusing (Deanon, 1988; Wang et al., 1998). Empig et al. (1962) report some resistance in Philippine native material, such as „Los Baños native‟ which has possibly been used in the local breeding research. J. Acosta8 conducted research on the inheritance of tomato bacterial wilt resistance at the University of Hawaii (Acosta et al., 1964) and one can hypothesize (i) that he took Philippine material to Hawaii9 and conversely (ii) that he brought material back home (to the Philippines). North Carolina material entered Philippines breeding program according to Mew & Ho (1977). These authors, on the basis of a personal communication of J.R. Deanon, indicate that „Venus‟ (North Carolina line) entered the pedigree of the Philippine line „UPCA1169‟, together with a „CA64-1169‟ of un-mentioned origin (Figure 1). „UPCA1169‟ is itself at the origin of other Philippines material such as „VC8-1-2‟ and „VC9-1‟ (Mew & Ho, 1977; Wang et al., 1998; Scott et al., 2005). The origin(s) of the resistance of other valuable Philippine material, such as „TML46‟, „TML114‟, „R3034‟, or „HSBW‟ mentioned in the Hawaii section is not known.

6 Dr H. Fukuoka, National Institute of Vegetable and Tea Science, Kusawa 360, Ano, Tsu, Mie 514-2392,

Japan. 7 „BF-Okitsu 101‟ was obtained by A. Kotani, T. Kuriyama, H. Shimada-Mochizuki, S. Sakuma, and I.

Suzuki (H. Fukuoka, pers. com.). 8 According to http://www.tofil.ph/awardee_profile.php?id=78, J Acosta won a Rockefeller scholarship to

the University of Hawaii in 1958. This is also indicated in Acosta (1963). 9 This hypothesis is consistent with the information of Gilbert to Laterrot (Laterrot et al., 1978) that a

Philippine source had been used in Hawaii and is at the origin of „Hawaii 7996‟.

12

French West Indies (INRA & IRAT) and Burkina Faso (IRAT) In the French West Indies, the research on tomato bacterial wilt resistance started in

1963 at IRAT, and in 1964 at INRA, with some collaboration between the two institutes. According to Cordeil & Digat (1967) a collection from Rio Pedras station of the

University of Puerto Rico was introduced at IRAT Guadeloupe beginning of 1964. These authors mention a variable tolerance to bacterial wilt of some lines such as „199 PR‟, „Platillo 78‟ and L. pimpinellifolium under local conditions. „199 UPR‟, a derivative from the double cross made at Puerto Rico University [„Platillo‟ X S. pimpinellifolium] X [a tomato of unknown identity X „Platillo‟] -see above-, was chosen for its good tolerance to bacterial wilt (Daly, 1976). After pedigree selection, the F8 lines „199 UPR -39.15‟ and „199 UPR -39.16‟ were obtained by Daly (1976). These lines were described as having small watery and not fleshy fruits, with greenish gel around the seeds, and displaying many concentric cracks (IRAT, 1965; Cordeil & Digat, 1967). Both were crossed with „Floralou‟, a variety of good quantitative and qualitative yield in the conditions of the French West Indies (Cordeil & Digat, 1967; Digat & Derieux, 1968; Daly, 1976). After pedigree selection, the lines „IRAT L3‟ (Daly, 1976; Laterrot et al., 1978) -Fig.6- and „Farako-Ba‟ (D‟Arondel de Haye, unpubl., IRAT, 1974, 1975; Laterrot et al., 1978; Rouamba et al., 1988) were respectively obtained in Martinique and Burkina Faso.

The INRA program in Guadeloupe was based on the use of „CRA66‟. The origin of this line is still controversial. It is given by Digat & Derieux (1968) and Anaïs (pers. com.) as one of the many small fruited tomato ecotypes grown in Guadeloupe at that time, and known there as „tomadoses‟. Digat & Derieux described „CRA66‟ as a vigorous line bearing small, pink, and bitter fruits with resistance to bacterial wilt. However another origin of CRA66 is suggested by Kaan (pers. com.) as being „OTB2‟, because the phenotype of „CRA66‟ is very different from the phenotype of the tomadoses: the plant is more vigorous, the leaves have a spreading leaf growth habit, the flowers display an exerted style, the fruits are fasciated and although of pink colour, they have a larger size than tomadoses fruits. There is a green gel inside the fruit, the taste of which is more acrid and extremely bitter, and its bacterial wilt resistance level is higher. However, Suzuki et al. (1964) reported „OTB1‟ was pink fruited, whereas they described „OTB2‟ as red fruited (but segregating for several traits). As IRAT experimented in Guadeloupe „OTB1‟, „OTB2‟ and many other bacterial wilt resistant lines in the mid- 1960s (IRAT 1964, 1965), it is plausible that valuable material „reached‟ INRA by some path and perhaps under a distorted identity for some reason. If „CRA66‟ = „OTB2‟, then all the French West Indies material would derive indirectly from North Carolina material -see Fig. 1-. Comparison of molecular fingerprinting of „OTB2‟ and „CRA66‟ is necessary for elucidating the identity of these two lines.

Crosses started in the 1960s between „CRA66‟ and the susceptible commercial type „Floradel‟ resulted in the varieties „Cranita‟ (Fig.7), „CRA74‟ & „Carette‟ (Fig. 8 & 9), and „Caraibo‟ (= „Caraibe‟) -Fig. 9- that were respectively released in 1971, 1973, 1975, and 1980 (Anaïs, 1986, Anaïs 1997). „CRA84-26-3‟ and „Caravel‟ are offspring of the cross [„Caraibo‟ X „HC8‟] where „HC8‟ is a heat-tolerant and bacterial wilt resistant line derived from the cross „Hawaii 7996‟ x „Campbell 28‟. Later on „Caraibo‟, „HC8‟ and „Caravel‟ were used as genitors in a recurrent selection for recombining their bacterial wilt

13

resistance and agro-climatic adaptation to French West Indies conditions, together with resistance to Begomoviruses (Ano et al., 2002; Ano et al., 2004).

Taiwan AVRDC started tomato breeding in 1972, and from 1973-1980 emphasized the

development of breeding lines with heat-tolerance and bacterial wilt resistance (Opena et al., 1989). Sources of BW resistance frequently used as parents in AVRDC breeding included varieties „Venus‟ and „Saturn‟ from North Carolina State University and lines „VC 11-3-1-8‟, „VC 8-1-2-7‟, „VC 48-1‟ from the University of the Philippines. Most AVRDC bacterial wilt resistant lines developed in the 1970‟s such as „CL8d-0-7-1‟ (derived from „VC11-1-2-1B‟ x „Venus‟), „CL9-0-0-1-3‟ (derived from „VC11-1-2-1B‟ x „Saturn‟), and „CL11d-0-2-1‟ (derived from „VC9-1-2-9B‟ x „Venus‟) were bred from crosses of these two sources. Many AVRDC lines developed in the late 1970‟s and early 1980‟s such as „CL1131‟, „CL5915‟ (Fig.10), and „CLN65‟ (Fig. 11) arose from complex crosses involving North Carolina, Philippines, or AVRDC lines with BW resistance derived from the above sources. High levels of BW resistance were detected in „L285‟, a small-fruited S. lycopersicum germplasm accession from Taiwan (Chang #1) but this source was not used at AVRDC in breeding because it was thought that its bacterial wilt resistance and small fruit size were closely associated (Opena et al., 1992). In 1985 AVRDC received resistant lines developed in Guadeloupe, including „CRA84-58-1‟ and „CRA84-26-3‟, that combined BW resistance and large fruit size. CRA lines were crossed to heat tolerant and BW resistant AVRDC lines which led to the development of AVRDC lines „CLN1462‟, „CLN1463‟ (Fig.12), „CLN1621‟, „CLN2026‟, and many others.

2. Limit of the reliability of the survey: insufficient accuracy of the information The information found in the literature is often vague and sometimes inconsistent

between publications. The absence of published pedigrees for many of the important varieties resistant to bacterial wilt is a real impediment for ascertaining the original sources of their resistance. Henceforth, Figure 1 is the result of our interpretation of sometimes blurred information as exemplified below.

Names of the accessions Depending on the publications, the accessions used in the various trials or breeding

programs are not exactly named the same way. This is due in some cases to probable renaming such as for „199 PR‟ (Cordeil & Digat, 1967), which is also found as “199” (Digat & Derieux, 1968), „199 UPR‟ (Daly, 1976), and „UPR 199‟ (Kaan et al., 1969). The same situation is encountered for „P.I. No. 3814‟ (Kelman, 1953), also found as „Beltsville #3814‟ (Henderson & Jenkins, 1972a; Hanson et al., 1998), „Beltsville No. 3814‟ (Henderson & Jenkins, 1972b), and „Beltsville 3814‟ (Laterrot et al., 1978). However other variations of names such as for „H7997‟ also found as „H7997S‟ and „H7997L‟, „H7998‟ also found as „H7998S‟ and „H7998M‟, or „CRA66‟ found as „CRA66P‟ and „CRA66S‟ in Wang et al. (1998) and Scott et al. (2005) do not mean further selections but encode only the name of the person who provided the seeds used in the trials. These latter authors mention „BF-Okitsu‟, otherwise found as „BF-Okitsu 101‟ in Laterrot (1999).

14

Despite the word “Okitsu” (location in Japan where breeding was carried out, H. Fukuoka, pers. com.) is present in the names of bacterial wilt resistant „BF-Okitsu 101‟ and of bacterial canker resistant „Okitsu Sozai n°110 (Kuriyama & Kuniyasu, 1974), these accessions should not be confused with each other.

Contradictory data found in the literature Apart from the controversy concerning the identity of „CRA66‟, other contradictory

information is found in the literature. Particularly in the case of „Hawaii 7996‟and the other Hawaii 79## accessions, resistance was said to originate from a Philippines accession according to several personal communications between breeders in the 1970s, but according to Acosta et al. (1964) „PI 127805A‟ and North Carolina material are at the origin of the resistance to bacterial wilt of Hawaiian material. It is not possible to reconcile these conflicting statements other than to say that the various sources of resistance were introduced to the Hawaiian program over time. Therefore we left all options on Figure 1.

Unclear origin of some accessions We found no original information on the origin or pedigree of „Beltsville 3814‟ and

further search of 1960s and 1970s publications of Beltsville USDA research station is needed. Another case concerns the unclear relationship between the Peruvian S. pimpinellifolium „PI 127805‟ collected in 1938 and maintained by the USDA Northeast Regional PI Station and „PI 127805A‟ obtained in 1953 and field selected for resistance through 9 generations in Hawaii according to Acosta et al. (1964). Furthermore, „5808-2‟ was derived from „PI 127805A‟ according to Mohanakumaran et al. (1967) but is given as an inbred line of an anonymous L. pimpinellifolium according to Acosta (1963).

Likeness between accessions Phenotypically „BF-Okitsu‟ is very close to „Hawaii 7998‟ (J. Scott and J. Wang, pers.

obs.), though the published information (Fig.1) does not indicate a closer relationship than the presence of North Carolina material in both their pedigrees. Comparison of their molecular fingerprinting would be worthwhile to clarify their genetic relationship.

3. Sources of resistance and inheritance patterns The global survey of the major breeding research for tomato bacterial wilt resistance

points out that the main sources of resistance used worldwide are perhaps only half a dozen accessions of S. pimpinellifolium („PI 127805A‟), S. lycopersicum var. cerasiforme („PI 129080‟), S. lycopersicum var. pyriforme („Beltsville 3814‟), a progeny from a cross between a S. pimpinellifolium and S. lycopersicum („199 UPR‟), S. lycopersicum („Mulua‟) and the enigmatic Philippine accession used in Hawaii in the 1970s. This number could be extended to seven accessions if one adds „CRA66‟ by assuming it is a Guadeloupe tomadose and not a progeny from „OTB2‟. Whether there are 6 or 7 main sources of resistance, the genetic basis of tomato resistance mechanisms used worldwide for breeding is quite narrow. North Carolina material has been integrated in most of the other breeding programs, in particular those of Hawaii, Japan, Philippines and Taiwan. By combining NC material, or not, with other sources of

10 „Okitsu Sozai n°1‟ resistance to Clavibacter michiganensis (formerly named Corynebacterium michiganense) originates from S. habrochaites (L. hirsutum var.

glabratum) „PI 134418‟.

15

resistance, and breeding in geographical areas where different strains of bacterial wilt are prevalent, the breeders exploited the genetic potentialities at their disposal, and created material resisting a wide range of bacterial wilt strains as exemplified by the results obtained in the worldwide trial carried out by Wang et al. (1998). Indeed, the top nine resistant accessions which had high levels of resistance in almost all 12 locations tested (>90% survival on average) were developed in Hawaii („H7996‟, „H7997 S and L‟, „H7998 S and M‟), Philippines („TML46‟ and „TML114‟, „R3034‟), and Japan („BF-Okitsu‟).

Other sources of resistance in wild tomatoes have been described sporadically in

the literature in accessions of the same species (S. pimpinellifolium, cherry and pear S. lycopersicum) as well as in other wild relatives of tomato (Laterrot & Kaan, 1977; Jaworski et al., 1987; Anaïs, 1997; Mohamed et al., 1997; Carmeille et al. 2006b; Hai et al., 2008). From these results, it seems that resistance to bacterial wilt is not that frequent in tomato germplasm. The high genetic diversity displayed by Ralstonia solanacearum complex (Fegan & Prior, 2005) and the strong interactions between strains and resistant material (Lebeau et al., 2011) suggest that various resistance mechanisms, including strain specific ones, exist in tomato resistant germplasm. Therefore, breeders have some opportunities at their disposal to enlarge the relatively narrow range of resistance sources primarily used so far, and to continue accumulating different mechanisms of resistance in tomato genotypes to obtain better stability of resistance in different environments. However, they might be limited by the fact that some bacterial strains are not controlled by any resistant accession (see section 5. below).

Inheritance studies have focused mostly on F2, F3 and RILs progenies of „Hawaii

7996‟ crossed with the susceptible „WVa700‟ (S. pimpinellifolium). Several QTLs of resistance have been identified, including a major QTL on chromosome 6 effective towards „GMI8217‟, an isolate of race 1 biovar 1 (Thoquet et al. 1996a & b); of „Pss4‟, an isolate of race 1, biovar 3, phylotype 1 (Wang et al., 1998); and „JT516‟, an isolate representative of race 3-phylotype II (Carmeille et al., 2006a). Wang et al. (2000) identified another major QTL of resistance of „Hawaii 7996‟ effective towards „Pss4‟ and located on chromosome 12. Several minor QTLs located on chromosomes 3, 4, 8, some of which having a season dependent expression (Carmeille et al., 2006a) have also been identified. Mejia et al (2009) confirmed the QTLs on chromosome 6 and 12 were associated with resistance in „Hawaii 7996‟ observed in Guatemala field evaluation against local phylotype I strains. Work is ongoing at AVRDC for adding markers to the QTLs regions of „Hawaii 7996‟ associated to resistance to bacterial strains belonging to phylotype I, in order to develop tools for marker assisted selection.

QTLs of resistance of the resistant line „L285‟ effective towards „UW364‟ an isolate of race 1, biovar 4, have also been located on chromosomes 6, as well as on chromosomes 7 and 10 (Danesh et al., 1994). Pattern of resistance derived from CRA66 has been described as polygenic (Prior et al., 1994) but no molecular data are available for this source.

16

4. Resistance to bacterial wilt is sometimes associated to resistance to other

bacterial and fungal pathogens Kaan & Laterrot (1977) were the first to mention a quantitative resistance to

Fusarium wilt (Fusarium oxysporum f.sp. lycopersici) race 2 in lines bred for bacterial wilt resistance in Puerto Rico, North Carolina, and the French West Indies and they suggested this relation to be more likely of a pleiotropic nature than being due to a genetic linkage. Further, Laterrot & Kaan (1978) as well as Laterrot et al. (1978) have pointed out the frequent association of both these partial resistances with the partial resistance to a third vascular disease, i.e. bacterial canker caused by Clavibacter michiganensis. These authors exemplified this relation between the resistance to the three diseases on a set of varieties bred for bacterial wilt in North Carolina („NC 72 TR 4-4‟, „MR4‟, „Venus‟ and „Saturn‟), in the French West Indies („Carette‟, „53 RC‟, „IRAT L3‟), in Burkina Faso („Farako-Ba‟), and in Hawaii („Hawaii 7996‟). They checked that the resistance to Fusarium wilt race 2 observed was not due to the gene I-2. „Kewalo‟ was the only variety resistant to bacterial wilt that they found susceptible to bacterial canker, and of a low level of resistance to Fusarium wilt race 2. This exception suggests that the mechanisms controlling the resistance to bacterial wilt can be dissociated, in some genotypes, from those involved in the resistance to the two other vascular diseases. Unfortunately Laterrot & Kaan (1978) and Laterrot et al. (1978) did not test bacterial wilt resistant material created in the Philippines and Taiwan, and the general picture of the relationships between the resistance to bacterial wilt and the two other vascular diseases is incomplete.

The reciprocal relationship between the resistance to bacterial canker and the two other diseases is verified in some cases, as for instance for „Okitsu Sozai n°1‟ whose resistance to bacterial canker originates from S. hirsutum var. glabratum PI 134418. This line is also partially resistant to Fusarium wilt race 2 (Laterrot, unpub. results) as well as to some strains of Ralstonia solanacearum (Lebeau et al., 2011). But „Plovdiv 8-12‟, the resistance of which to bacterial canker originates from a S. pimpinellifolium, is susceptible to Fusarium wilt race 2 (Laterrot et al. 1978); its behaviour towards bacterial wilt is unknown.

The bacterial wilt resistant „Hawaii 7998‟ was resistant to a race of bacterial spot

later confirmed to be race T1 (Scott & Jones 1986). Later, Scott et al (1995b) discovered „Hawaii 7981‟, a line susceptible to race T1, was resistant to bacterial spot race T3 while „Hawaii 7998‟ was susceptible. Thus, bacterial spot resistance has been found in bacterial wilt resistant lines from Hawaii. For race T1, resistance to bacterial wilt was not correlated with bacterial spot resistance in an F2 population suggesting separate genes were responsible for resistance to each disease (Scott et al., 1988). Resistance to bacterial spot race T4 has been seen in some bacterial wilt resistant breeding lines derived from „Hawaii 7997‟ even though this line is not resistant to race T4 (Scott et al., 2010). The genetic control of this response is not known but again illustrates the association of bacterial wilt resistance with resistance to another disease. A further example is that „Hawaii 7998‟ had resistance to bacterial canker (Panthee and Gardner, 2010) especially the foliar phase. These examples and those mentioned before indicate the existence of frequent associations between the resistance to some

17

bacteria (vascular or not) and even fungi (vascular) within single tomato genotypes. Hence, Scott (1997) advised that when searching for resistance to a given bacterial pathogen, breeders should not overlook genotypes resistant to other bacteria. In this regard, resistance genes can emerge by combining genotypes with some reported bacterial resistance even when one of the parent lines does not show resistance to a particular disease/race. For instance, Hutton et al. (2010) reported that an allele from „Hawaii 7998‟ was associated with resistance to bacterial spot race T4 even though this line was susceptible to race T4, suggesting epistasis with genes from the other parent that was T4 resistant. Another possibility is that the pleiotropic nature of resistance in tomato against several diseases could be associated with higher level of expression of systemic acquired resistance (SAR). Lin et al (2004) found over-expression of Arabidopsis NPR1 (non-expresser of PR genes) gene in a susceptible tomato line could enhance resistance to both Fusarium wilt race 2, bacterial wilt, as well as bacterial spot and gray leaf spot. In this study, they did not test the enhanced resistance against the bacterial canker pathogen.

Lastly, Rouamba et al. (1988) tested material resistant to bacterial wilt towards a

fourth vascular disease, the Verticillium wilt caused by Verticillium race 2, but they found only a loose relationship between both resistances since only two („IRAT L3‟ and „Farako-Ba‟) out of ten lines tested, were resistant to both diseases.

5. Grafting of susceptible cultivars on resistant rootstocks, an alternative to resistant cultivars

Given the difficulty to create highly resistant lines with good commercial quality, grafting susceptible scions on resistant rootstocks remains an alternative to the cultivation of resistant cultivars. As early as 1969, Gilbert and Chin pointed out that highly resistant tomato lines with poor fruit quality, could be efficiently used as rootstocks on which susceptible scions of good fruit quality could be grafted. These authors reported the bacterial wilt resistance of the root system as being effective even when completely susceptible scions are used. This technique is still used nowadays (Cardoso et al., 2006; Wang et al., 2009), though the protection provided by the rootstock is sometimes incomplete (Nakaho et al., 2004). Indeed, tomato resistant material harbours the bacteria symptomlessly and the resistance is associated with the ability of the plant to restrict bacteria invasiveness (Grimault et al., 1993). The absence of incompatible interactions in tomato resistant lines (no symptoms, no latent infection) has been confirmed by Lebeau et al. (2011) by testing a core collection of bacterial wilt resistant accessions with a core collection of bacterial strains.

Eggplant is an alternative rootstock for cultivating susceptible tomatoes in contaminated conditions. It was shown to provide a better protection than tomato rootstocks (AVRDC, 1998). This result was confirmed and extended by Lebeau et al. (2011) who found that apart from common cases of latent infection for some eggplant accessions and all tomato accessions, there also exist incompatible interactions between some eggplant resistant lines and some bacterial strains. Further, some eggplant lines control bacterial strains that are not controlled by any of the tomato resistant lines tested so far, as exemplified by Carmeille et al. (2006b), Wicker et al. (2007) and Lebeau et al. (2011).

18

Conclusion The earliest breeding efforts for tomato resistance to bacterial wilt started in Puerto

Rico and two American States (North Carolina, Hawaii) 80 years ago followed by programs in the 1950s in Japan and the Philippines. The programs carried out by French research institutes in the Caribbean (and Burkina Faso) started during the course of the 1960s, whereas AVRDC started at the beginning of the 1970s. There were many exchanges of material between the breeding programs for bacterial wilt resistance carried out in several US States, and between them and the Philippines. AVRDC is using and recombining now the resistances bred in the USA, the Philippines and the French West Indies. The breeding material created in Japan was mostly used locally, and perhaps also in Guadeloupe, if one assumes that the Japanese line „OTB2‟ is equivalent to „CRA66‟, which is not certain.

Our attempt to draw a general picture of the main sources of resistance to bacterial wilt, of the main breeding programs for this resistance, of their most frequently mentioned resistant varieties, and of the relationships between varieties, is based on a careful work of assembling bits and pieces dispersed in many publications. Given the unavailability of complete information, it is not now possible to come up with a better picture than the one we present here, though more information can emerge out of archives of the scientists, Universities and research institutes involved. Henceforth the synthesis provided here displays the most probable general picture, but it may include some mistakes in the absence of further information.

We added to this survey complementary information on some peculiarities of tomato bacterial wilt resistance. The frequent association of bacterial wilt resistance with resistance to other bacterial or fungal diseases should be of strong interest for breeding and/or research on its genetic basis. Breeding over many decades succeeded in eliminating a number of undesirable traits initially associated to high level of bacterial wilt resistance, but breeding efforts are still ongoing for obtaining large fruit size in resistant material. The adaptation of the breeding material to hot environmental conditions is often mentioned as necessary for obtaining breeding lines which are resistant and agronomically acceptable.

On the whole, we hope this paper to be useful for further research using the plant material mentioned, in particular for comparative genetic studies and breeding concerning the resistance of tomato to bacterial wilt and to other vascular diseases. Acknowledgments The authors are very grateful to Ph. Prior and E. Wicker (phyto-bacteriologists at CIRAD La Réunion Island, Mascarenes) for having motivated the authors to write this historical review, to F. Kaan and G. Anaïs (retirees of INRA Guadeloupe) for having provided precious complements of information, to H. Fukuoka (National Institute of Vegetables & Tea Science, Tsu, Mie, Japan) for having fully clarified the relationships between former Japanese scientists, former Japanese Institutes and tomato lines and for having translated key passages of Suzuki et al. (1964). We thank also the consortium of private companies (Vilmorin, Gautier Semences, DeRuiter Seeds, Enza Zaden, Nunhems, Rijk Zwaan) who financed (2007-2010) research based on the present review. Last but not least, we acknowledge Ch. Olivier (librarian of INRA GAFL, Montfavet, France), M.L.

19

Abinne (librarian at INRA-CRAAG, Guadeloupe),and the librarians at North Carolina State University, the University of Hawaii, AVRDC, and the University of Puerto Rico along with Linda Wessel-Beaver for having provided reprints of numerous archives. Acronyms found in some names of tomato lines or in related literature UPR: University of Puerto Rico NCSU: North Carolina State University UPLB = UPCA: University of Philippines Los Banos = University of Philippines College of Agriculture CRA: Centre de Recherche Agronomique des Antilles (INRA)

20

Figure 1: General scheme of the relationships between worldwide programs and lines. Notations in color correspond to Literature Cited starting on page 23.

21

Table 1. Summing up of the phenotype of some breeding lines created or used in the breeding programmes of North Carolina, Hawaii, Florida, Japan, Philippines, French West Indies, and Taiwan.

origin

line or accession name growth habit fruit shape fruit size

fruit colour source

North Carolina University NC 72 TR 4-4 indeterminate

slightly flattened 60-100 g red

Laterrot et al. (1978); INRA germplasm database

North Carolina University MR4 indeterminate

Laterrot et al. (1978)

North Carolina University NC1953-60N 6,5 g

Suzuki et al. (1964)

North Carolina University NC19/53-64N 7,6 g

Suzuki et al. (1964)

North Carolina University Saturn indeterminate deep globe 100-140 g red

Kaan et al. (1975); INRA germplasm database; Henderson & Jenkins (1972)

North Carolina University Venus indeterminate slightly oblate > 180 g red

Laterrot et al. (1978); INRA germplasm database; Henderson & Jenkins (1972)

Hawaii University HES 5808-2 indeterminate 15g

Hawaii University H7996 determinate small oblate 20-60g red

INRA germplasm database, Scott. pers. com.

Hawaii University H7997 indeterminate small oblate 30-80 g red

Wang et al. (1998), Scott et al. (2005), Scott. pers. com.

Hawaii University H7998 indeterminate small oblate 30 g red

Wang et al. (1998), Scott et al. (2005), Scott. pers. com.

Hawaii University Kewalo determinate flattened 140-180 g red

INRA germplasm database

University of Florida Neptune determinate 123-136g red

Scott et al. (1995a)

Hort. Res. Sta.,Japan BF-Okitsu indeterminate 15-20 g red

Wang et al. (1998)

Hort. Res. Sta.,Japan OTB1 13,8 g pink

Suzuki et al. (1964)

Hort. Res. Sta.,Japan OTB2 30,6 g red

Suzuki et al. (1964)

22

Philippines University UPCA1169 determinate 20-30 g Source ?

Philippines University CA-64-1169

Philippines University VC8-1-2

Philippines University VC9-1

Philippines University TML46 determinate oblate/oblong 30 g red/pink

Wang et al. (1998)

Philippines University TML114 determinate oblate/oblong 40 g red/pink

Wang et al. (1998)

Philippines University R3034

semi determinate deep oblate 30-60 g red

Wang et al. (1998)

INRA, Guadeloupe CRA66 indeterminate

slightly flattened 20-60 g pink

INRA germplasm database. This line is recorded as red fruited in Wang et al. (1998); Scott et al. (2005)

INRA, Guadeloupe Cranita indeterminate pink

Messiaen et al.(1978); Laterrot

INRA, Guadeloupe CRA74 indeterminate wide, deep medium

Kaan et al. (1975)

INRA, Guadeloupe Carette indeterminate

slightly flattened 100-140 g red

Laterrot et al. (1978); INRA germplasm database

INRA, Guadeloupe

Caraibo = Caraibe determinate

flattened / oblate

140-180 g / 150 g red

Anais (1986); Ano et al.(2004); INRA germplasm database

INRA, Guadeloupe Caravel determinate oblate 150-300 g red

Wang et al. (1998)

INRA, Guadeloupe CRA84-26-3 determinate

Hanson et al. (1996)

IRAT (Guadeloupe & Martinique) IRAT L3 indeterminate round 45 g red

(Laterrot et al., 1978); INRA germplasm database; Denoyés (1988)

IRAT (Burkina Faso) Farako-Ba indeterminate round 140-180 g red

INRA germplasm database

AVRDC, Taiwan CL5915 determinate oblong 50 g red

Wang et al. (1998)

AVRDC, Taiwan CLN65 determinate oblate 70 g red

Wang et al. (1998)

AVRDC, Taiwan CLN1463 indeterminate globe 150-200 g red

Wang et al. (1998)

AVRDC, Taiwan L285 indeterminate plum 30 g red

Wang et al. (1998)

23

Literature cited

1. Acosta J.C., 1963. Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill. PhD Thesis, University of Hawaii (Agriculture, Plant Pathology). On line accessible at: http://scholarspace.manoa.hawaii.edu/bitstream/10125/11673/2/uhm_phd_6402645_r.pdf

2. Acosta, J., Gilbert, J., and Quinon, V.L., 1964. Heritability of bacterial wilt resistance in tomato. Proceedings of the American Society for Horticultural Science 84:455-461.

3. Anaïs G., 1986. Utilisation de la résistance variétale dans la lutte contre le flétrissement bactérien de la tomate Pseudomonas solanacearum E.F. Smith. Bull. Tech. D‟Information 409/411 : 449-452.

4. Anaïs G., 1997. La tomate. Pp 591-605 In : L‟Amélioration des Plantes tropicales. A. Charrier, M. Jacquot, S. Hamon, D. Nicolas (Sc. Eds), Ed. CIRAD, ORSTOM..

5. Anais G., pers. com. 2010. 6. Ano G., Anaïs G., Marival P., 2002. Création de variétés de tomate résistantes à

Ralstonia solanacearum et PYMV, adaptées aux régions tropicales. Proceedings of the XXXVIIIth Meeting of the Carribean Food Crops Society, Martinique : 234-238.

7. Ano G., Anaïs G., Marival P., Chidiac A., 2004. L‟amélioration de la tomate pour les régions tropicales de plaine: travaux en Guadeloupe. Les familles CRAPY associent les résistances aux Bégomovirus PYMV et TYLCV à la résistance au Ralstonia solanacearum (race 1). Phytoma, la défense des végétaux 573 : 23-25.

8. AVRDC Report 1998. Asian Vegetables Research and Development Center, Taiwan.

9. Azzam H., 1964. Tomato breeding for the tropics. Proc. of the 2nd Annual Meeting of the Caribbean Food Crops Society, Bridgetown, Barbados 2: 56-59.

10. Balatero, C. H., and Hautea, D. M. 2001. Identification of AFLP and RGA markers associated with bacterial wilt resistance QTL derived from tomato Lycopersicon pimpinellifolium. p.225-243 in Solanaceae V: advances in taxonomy and utilization, edited by R. G. van den Berg, G. W. M. Barendse, G. M. van der Weerden and C. Mariani. Nijmegen University Press.

11. Balatero C.H., Hautea D.M., Narciso J.O., Hanson P.M., 2005. QTL mapping for bacterial wilt resistance in Hawaii 7996 using AFLP, RGA and SSR markers. Pp 301-308 In: Bacterial wilt: the disease and the Ralstonia solanacearum complex. C. Allen, P. Prior, A.C. Hayward (Sc. Eds), American Phytopathological Society (APS), St Paul, USA.

12. Barham W.S. Ellis D.E., 1951. Sources of resistance to late blight and bacterial wilt. Rept. Tomato Genet. Coop. 1: 2.

13. Borchers E.A.,Nevin C.S., 1954. Quantitative estimation of a bitter principle in tomato. Proc. Amer. Soc. Hort. Sci. 63:420-426.

14. Cardoso, S. C.; Soares, A. C. F.; Brito, A. dos S.; Carvalho, L. A. de; Ledo, C. A. da S., 2006. Potential of Hawaii 7996 hybrid as rootstock for tomato cultivars. Bragantia 65(1): 89-96.

24

15. Carmeille, A., Caranta, C., Dintinger, J., Prior, P., Luisetti, J., and Besse, P. 2006a. Identification of QTLs for Ralstonia solanacearum race 3-phylotype II resistance in tomato. Theoretical and Applied Genetics 113 (1):110-121.

16. Carmeille A., Prior P., Kodja H., Chiroleu F., Luisetti J., Besse P., 2006b. Evaluation of resistance to race 3, Biovar 2 of Ralstonia solanacearum in tomato germplasm. J. Phytopathology 154: 398-402.

17. Celine V.A., Chandrmony D., Gokulapalan C., Rajamony L., 2003. Heat tolerance and bacterial wilt resistance of tomato genotypes in the humid tropics of Kerala. Rept. Tomato Genet. Coop. 53: 11:13.

18. Cook M.T., 1934. Annual report of the division of botany and plant pathology (tomatoes). In Puerto Rico Univ. Agric. Expt. Sta. Ann. Rpt. (1933/1934): 131-132.

19. Cook M.T., 1935. Annual report of the division of botany and plant pathology (wilt disease of tomato). In Puerto Rico Univ. Agric. Expt. Sta. Ann. Rpt. (1934/1935): 25-26.

20. Cordeil J., Digat B., 1967. Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française. Proc. of the 5th Annual Meeting of the Caribbean Food Crop Society, Paramarimbo, Surinam, 5: 91-98.

21. Daly, P. 1976. "IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies. Agronomie Tropicale 31 (4):398-402.

22. Danesh D., Aarons S., Mcgill G.E., Young N.D., 1994. Genetic dissection of oligogenic resistance to bacterial wilt in tomato. Molecular Plant Microbe Interaction 7: 464-471.

23. Daunay M.C., 1977. Une résistance de la tomate à trois maladies vasculaires. Mémoire de fin d‟Etudes, Ecole Nationale des Ingénieurs des Techniques Horticoles, Angers, FRA. Ed. INRA. 72 pp + annexes.

24. Deanon, J. R. Jr. 1988. Biochemical bases of screening tomato for bacterial wilt resistance. p.83-99 In: Current problems on fruits and vegetables. Philippines Council for Agriculture Forestry and Natural Resources Research and Development, Los Banos, Laguna, Philippines.

25. Deanon, Pers. com to Mew & Ho, 1977. 26. Deberdt P., Olivier J., Thoquet P., Grimsley N., Prior P., 1999a. Quantitative

resistance loci to bacterial wilt located on tomato chromosome 6 in near isogenic lines for the Mi gene. Proc. of the First Australasian Soilborne Disease Symp., R.C. Magarey (Ed), Bureau of Sugar Experiment Stations, Brisbane, Australia: 93-94.

27. Deberdt, P., Quénéhervé, P., Darrasse, A., and Prior, P. 1999b. Increased susceptibility to bacterial wilt in tomatoes by nematode galling and the role of the Mi gene in resistance to nematodes and bacterial wilt. Plant Pathology 48:408-414.

28. Digat, B., and M., Derieux. 1968. A study of the varietal resistance to bacterial wilt. p.95-101 in Proceedings of the Caribbean Food Crops Society, 6th Annual meeting, St Augustine, Trinidad.

29. Empig L.T., Calub A.G., Katigbak M.M., Deanon J.R., 1962. Screening tomato, eggplant, and pepper varieties and strains for bacterial wilt (P. solanacearum E.F.S.) resistance. Philippine Agriculturist (46): 303-314.

30. Fegan M., Prior P., 2005. How complex is the “Ralstonia solanacearum species complex”. In: Bacterial Wilt: the Disease and the Ralstonia solanacearum species

25

complex. C. Allen, P. Prior and C. Hayward (Eds), APS Press, St. Paul, Minnesota, USA , 449-462.

31. Gilbert J.C., Chinn J.T., 1969. Bacterial wilt resistant combinations. Rept. Tomato Genet. Coop. 19: 11.

32. Gilbert J.C., Tanaka J.S., 1965. Horticultural refinement of multiple disease resistant tomatoes in Hawaii. Hawaii Farm Science 14(1): 4-6.

33. Gilbert J.C., Tanaka J.S., Takeda K.Y., 1974. „Kewalo‟ tomato. Hortscience 9: 481-482.

34. Grimault V., Schmit J., Prior P., 1993. Some characteristics involved in bacterial wilt (Pseudomonas solanacearum) resistance in tomato. p.112-119 In: Bacterial wilt. ACIAR proceedings No. 45, edited by G. L. Hartman and A. C. Hayward. ACIAR, Canberra.

35. Hai T.T.H., Esch E, Wang J.F., 2008. Resistance to Taiwanese race 1 strains of Ralstonia solanacearum in wild tomato germplasm. European Journal of Plant Pathology 122 (4): 471-479.

36. Hanson P., 2010. pers. com. 37. Hanson, P. M., Licardo, O., Hanudin, Wang, J. F., and Chen, J. T. 1998. Diallel

analysis of bacterial wilt resistance in tomato derived from different sources. Plant Disease 82:74-78.

38. Hanson, P. M., Wang, J. F., Licardo, O., Hanudin, Mah, S. Y., Hartman, G. L., Lin, Y. C., and Chen, J. T. 1996. Variable reactions of tomato lines to bacterial wilt evaluated at several locations in Southeast Asia. HortScience 31:143-146.

39. Henderson W.R., Jenkins S.F., 1972a. „Venus‟ and „Saturn‟, tomato varieties resistant to Southern bacterial wilt. Hortscience 7: 346.

40. Henderson W.R., Jenkins S.F., 1972b. „Venus‟ and „Saturn‟, two new tomato varieties combining desirable horticultural features with southern bacterial wilt resistance. Bulletin 444 of Agricultural Experiment Station, North Carolina State University (Raleigh), July 1972.

41. Hutton, Samuel F., Jay W. Scott, Yang, Wencai, Sim, Sung-Chur, Francis, David M., and Jones, Jeffrey B. 2010. Identification of QTL Associated with Resistance to Bacterial Spot Race T4 in Tomato. Theor. Appl. Genet. 121 (7):1275-1287.

42. IRAT, 1964. Annual Report, Guadeloupe Martinique. Ed. IRAT, p. 433. 43. IRAT, 1965. Annual Report, Guadeloupe. Ed. IRAT, p. 563. 44. IRAT, 1974. Annual Report, Martinique. Ed. IRAT, p.134. 45. IRAT, 1975. Annual Report, Martinique. Ed. IRAT, p.141. 46. Jaunet, T. X., and Wang, J. F. 1999. Variation in genotype and aggressiveness of

Ralstonia solanacearum race 1 isolated from tomato in Taiwan. Phytopathology 89 (4):320-327.

47. Jaworski C.A., Phatak S.C., Ghate S.R., Gitaitis R.D., Widrlechner M.P., 1987. GA 1565-2-4 BWT, GA 219-1-2 BWT, GA 1095-1-4 BWT, and GA 1405-1-2 BWT Bacterial Wilt Tolerant Tomato. HortScience 22(2): 324-325.

48. Kaan F., 1976. Mémoire sur les travaux réalises. CRAAG, Station d‟Amélioration des Plantes, Guadeloupe. Doc. Interne.

49. Kaan F., Beramis M., Messiaen, C.M., 1969. Recherche de variétés de tomates aux Antilles. Proceed. VIIth Ann. Meeting Caribb. Food Crops Society, 7 : 173-181.

26

50. Kaan F., Laterrot H., 1977. Mise en évidence de la relation entre des résistances de la tomate à deux maladies vasculaires: le flétrissement bactérien (Pseudomonas solanacearum E.F. Sm.) et la Fusariose pathotype 2 (Fusarium oxysporum f.sp. lycopersici (Sacc.) Snyd. & Hans). Ann. Amélior. Plantes 27(I) : 25-34.

51. Kaan F., Laterrot H., Anaïs G., 1975. Etude de 100 variétés de tomate en fonction de l‟adaptation climatique et de la résistance à sept maladies sévissant aux Antilles. Nouv. Agron. Antilles-Guyane, (1,2) : 123-138.

52. Kelman, A., 1953. The bacterial wilt caused by Pseudomonas solanacearum. A literature review and bibliography. North Carolina Agricultural Experiment Station, Tech. Bul. N°99: 194 pp.

53. Krausz J.P., Thurston H.D., 1975. Breakdown of resistance to Pseudomonas solanacearum in tomato. Phytopathology 65: 1272-1274.

54. Kuriyama T., & Kuniyasu K, 1974. Studies on the breeding of disease resistant tomato by interspecific hybridization. III. On the breeding of a new tomato resistant to bacterial canker caused by Corynebacterium michiganense. Bull. of the Veg. and Ornament. Crops Research Station A, 1: 93-107.

55. Laterrot, H., Kaan, J.F. 1978. Resistance to Corynebacterium michiganense of lines bred for resistance to Pseudomonas solanacearum. Report of the Tomato Genetics Cooperative 28.

56. Laterrot, H., Brand, R., Daunay, M. C. 1978. La résistance à Corynebacterium michiganense chez la tomate. Annales Amélioration des Plantes 28 (5):579-591.

57. Lebeau A., Daunay M.C., Frary A., Palloix A., Wang J.F., Dintinger J., Chiroleu F., Wicker E., Prior P., 2011. Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex. Phytopathology 101(1): 154-165.

58. Lin, W.-C., Liu, C.-F., Wu, J.-W., Cheng, M.-L., Lin, Y.-M., Yang, N.-S., Black, L., Green, S.K., Wang, J.-F., and Cheng, C.-P. 2004. Transgenic Tomato Plants Expressing the Arabidopsis NPR1 Gene Confer Enhanced Resistance to A Spectrum of Fungal and Bacterial Diseases. Transgenic Research 13:567-581.

59. Mangin, B., Thoquet, P., Olivier, J., and Grimsley, N. H. 1999. Temporal and multiple quantitative trait loci analyses of resistance to bacterial wilt in tomato permit the resolution of linked loci. Genetics 151:1165-1172.

60. Mejía, L., Garcia, B. E., Fulladolsa, A. C., Ewert, E. R., Wang, J.-F., Scott, J. W., Allen, C. and Maxwell, D. P. 2009. Evaluation of recombinant inbred lines for resistance to Ralstonia solanacearum in Guatemala and preliminary data on PCR-based tagging of introgressions associated with bacterial wilt-resistant Line, Hawaii 7996. Rept. Tomato Genet. Coop. 59:32-4.

61. Messiaen C.M., 1989. Environmental influences on the severity of tomato bacterial wilt in the French West Indies: Interactions with varietal resistance. Pp 235-238 In: Tomato and Pepper Production in the Tropics. Proceed. Internat. Symp. on Integrated Management Practices; S.K. Green (Sc. Ed.), T.D. Griggs & B.T. Mc Lean (Publ. Eds), AVRDC, Shanhua, Taiwan.

62. Messiaen, C. M., Laterrot, H., and Kaan, F. 1978. Cumulate resistances to Pseudomonas solanacearum and to Meloidogyne icognita with determinate growth in tomato. p.48-51 in Vegetables for the hot humid tropics. Mayaguez Institute of Tropical Agriculture, Puerto Rico

27

63. Mew T.W., Ho W.C., 1977. Effect of soil temperature on resistance of tomato cultivars to bacterial wilt. Phytopathology 67: 909-911.

64. Mohamed M.El.S., Umaharan P., Phelps R.H., 1997. Genetic nature of bacterial wilt resistance in tomato (Lycopersicon esculentum Mill.) accession LA 1421. Euphytica 96: 323-326.

65. Mohanakumaran N., Gilbert J.C., Young R.L., 1967. Bacterial wilt resistant tomato lines with unusually high content of the alkaloid tomatin. Rept. Tomato Genet. Coop. 17: 41.

66. Nakaho K., Inoue H., Takayama T., Miyagawa H., 2004. Distribution and multiplication of Ralstonia solanacearum in tomato plants with resistance derived from different origins. J. Gen. Plant Pathol. 70: 115-119.

67. Opena R.T., Hartman G.L., Chen J.T., Yang C.H. 1992. Breeding for bacterial wilt resistance in tropical tomato. p.44-50 in: Proceedings of the 3rd international conference on plant protection in the tropics. Malayasian Plant Protection Society, Kuala Lumpur.

68. Opena R.T., Green S.K., Talekar N.S., Chen J.T., 1989. Genetic improvement of tomato adaptability to the tropics: progress and future prospects. Pp 71-85 In: Tomato and Pepper Production in the Tropics. Proceedings of the International Symposium on Integrated Management Practices, S.K. Green (Sc. Ed.), T.D. Griggs & B.T. Mc Lean (Publ. Eds), AVRDC, Shanhua, Taiwan.

69. Panthee, D and Gardner R.G. 2010. Identification of useful sources of bacterial wilt resistance in tomato: A challenge. Proc. 25th Annual Tomato Disease Workshop. P. 18[Abstr.]

70. Peter, K. V., Gopalakrishnan, T. R., Rajan, S., and Sadhan Kumar, P. G. 1993. Breeding for resistance to bacterial wilt in tomato, eggplant and pepper. p.183-190 in Bacterial wilt. ACIAR proceedings No. 45, edited by G. L. Hartman and A. C. Hayward. ACIAR, Canberra.

71. Prior P., Grimault V., Schmit J., 1994. Resistance to bacterial wilt (Pseudomonas solanacearum) in tomato: present status and prospects. In: Bacterial Wilt, the disease and its causative agent Pseudomonas solanacearum, A.C. Hayward & G.L. Hartman (eds.), Wallingford (UK), CAB International : 209-223.

72. Roque A., 1935. Annual Report of the Assistant Pathologist (wilt disease of tomatoes). Puerto Rico Univ. Agr. Expt. Sta., Ann. Rpt. (1934-1935): 31-33.

73. Rouamba A., Laterrot H., Moretti A., 1988. A case of relation between resistances to Pseudomonas solanacearum and Verticillum pathotype 2. Rept. Tomato Genet. Coop. 38: p.43.

74. Schmidt R., 1936. Tomato breeding. N.C. Agri. Expt. Sta. Ann. Rep. 59: 68. 75. Schmidt R., 1937. Tomato breeding. N.C. Agri. Expt. Sta. Ann. Rep. 60: 55-56. 76. Scott, J.W. 1997. Tomato improvement for bacterial disease resistance for the

tropics: A contemporary basis and future prospects. In: Proc. First International Conference on Processing Tomato/First International Symposium on Tropical Tomato Diseases, Recife, Brazil, ASHS Press, Alexandria, VA USA. p. 117-123.

77. Scott, J.W., Jones J.B., 1986. Sources of resistance to bacterial spot (Xanthomonas campestris pv. vesicatoria (Doidge) Dye) in tomato. HortScience 21(2):304-306.

28

78. Scott, J.W., Jones J.B., Somodi G.C., 2003. Development of a large fruited tomato with a high level of resistance to bacterial wilt (Ralstonia solanacearum). Rept. Tomato Genet. Coop 53:36-37.

79. Scott, J.W., Jones J.B., Somodi G.C., Chellemi D.O., Olson S.M., 1995a. „Neptune‟, a heat-tolerant, bacterial wilt-tolerant tomato. HortScience 30(3):641-642.

80. Scott, J.W., Jones J.B., Somodi G.C., Stall R.E., 1995b. Screening tomato accessions for resistance to Xanthomonas campestris pv. vesicatoria, race T3. HortScience 30(3):579-581.

81. Scott, J.W., Jones J. B., Vallad G. E., 2010. Breeding for Bacterial Wilt Resistance in Tomato: The Struggle Continues. Proc. 25th Annual Tomato Disease Workshop. P.19[Abstr.]

82. Scott J.W., Somodi G.C., Jones J.B., 1988. Bacterial spot resistance is not associated with bacterial wilt resistance in tomato. Proc. Fla. State Hort. Soc. 101: 390-392.

83 Scott, J.W., G.E. Vallad, and J.B. Jones. 2009. High level of resistance to bacterial wilt (Ralstonia solanacearum) obtained in large-fruited tomato breeding lines derived from Hawaii 7997. Acta Horticulturae 808:269-274.

84. Scott, J.W., J.F. Wang, and P.M. Hanson. 2005. Breeding tomatoes for resistance to bacterial wilt, a global view. In: Proceeding of the First International Symposium on Tomato Diseases, Orlando, Florida, USA. Acta Hort. (ISHS) 695:161-172.

85. Sonoda R.M., Augustine J.J., Volin R.B., 1979. Bacterial wilt of tomato in Florida: history, status, and sources of resistance. Proc. Fla. State Hort. Sci. 92:100-102.

86. Suzuki, I., Sugahara, Y., Kotani, A., Todaka, S., and Shimada, H. 1964. Studies on breeding eggplants and tomatoes for resistance to bacterial wilt. I. Investigations on method of evaluating the resistance and on the source of resistance in eggplants and tomatoes. Bulletin-Horticultural Research Station Ministry of Agriculture and Forestry Series A 3:77-106.

87. Theis T., 1950. Tomato diseases. Puerto Rico Univ. Agr. Expt. Sta., Ann. Rpt. (1950): 9.

88. Thoquet, P., Olivier, J., Sperisen, C., Rogowsky, P., Laterrot, H., and Grimsley, N. 1996a. Quantitative trait Loci determining Resistance to Bacterial Wilt in Tomato Cultivar Hawaii7996. Molecular Plant-Microbe Interactions 9:826-836.

89. Thoquet, P., Olivier, J., Sperisen, C., Rogowsky, P., Prior, P., Anaïs, G., Mangin, B., Bazin, B., Nazer, R., and Grimsley, N. 1996b. Polygenic resistance of tomato plants to bacterial wilt in the French West Indies. Molecular Plant-Microbe Interactions 9:837-842.

90. Thurston H.D., 1976. Resistance to bacterial wilt (Pseudomonas solanacearum). Proc. 1rst Int. Conference and Worskshop on the ecology and control of bacterial wilt caused by Pseudomonas solanacearum. Raleigh, July 1976: 58-62.

91. Vaneechoutte M, Kämpfer P, De Baere T, Falsen E, Verschraegen G. 2004. Wautersia gen. nov., a novel genus accommodating the phylogenetic lineage including Ralstonia eutropha and related species, and proposal of Ralstonia [Pseudomonas] syzygii (Roberts et al. 1990) comb. nov. Int J Syst Evol Microbiol. 54(Pt 2):317-27.

92. Walter J.M., 1967. Hereditary resistance to disease in tomato. Annual Review of Phytopathology, vol.5: 131-162.

29

93. Wang, J.-F., P. Hanson, and J.A. Barnes. 1998. Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato. In: P. Prior, C. Allen and J. Elphinstone (eds.), Bacterial Wilt Disease: Molecular and Ecological Aspects, Springer-Verlag, Berlin 269-275.

94. Wang, J. F., Olivier, J., Thoquet, P., Mangin, B., Sauviac, L., and Grimsley, N. H. 2000. Resistance of tomato line Hawaii7996 to Ralstonia solanacearum Pss4 in Taiwan is controlled mainly by a major strain-specific locus. Molecular Plant-Microbe Interactions 13 (1):6-13.

95. Wang HanRong; Ru ShuiJiang; Wang LianPing; Fang Li; Ren HaiYing; Feng ZhongMen, 2009. Control of tomato bacterial wilt with grafting. Acta Agriculturae Zhejiangensis 21(3): 283-287.

96. Warmke H.E., Cruzado H.J., 1949. Tomato breeding. Puerto Rico Fed. Expt. Sta. Ann. Rpt. For 1949: 9.

97. Wicker E., Grassart L., Coranson-Beaudu R., Mian D., Guilbaud C., Fegan M., Prior P., 2007. Ralstonia solanacearum strains from Martinique (French West Indies) exhibiting a new pathogenic potential. Appl. & Environ. Microbiology 73(21): 6790-6801.

98. Winstead N.N., Kelman A., 1952. Inoculation techniques for evaluating resistance to Pseudomonas solanacearum. Phytopathology 42: 628-634.

99. Yabuuchi E, Kosako Y, Oyaizu H, Yano I, Hotta H, Hashimoto Y, Ezaki T, Arakawa M. 1992. Proposal of Burkholderia gen. nov. and transfer of seven species of the genus Pseudomonas homology group II to the new genus, with the type species Burkholderia cepacia (Palleroni and Holmes 1981) comb. nov. Microbiol Immunol.36(12):1251-75.

100.Yabuuchi E, Kosako Y, Yano I, Hotta H, Nishiuchi Y. 1995. Transfer of two Burkholderia and an Alcaligenes species to Ralstonia gen. Nov.: Proposal of Ralstonia pickettii (Ralston, Palleroni and Doudoroff 1973) comb. Nov., Ralstonia solanacearum (Smith 1896) comb. Nov. and Ralstonia eutropha (Davis 1969) comb. Nov.Microbiol Immunol. 39(11): 897-904.

101.Yarnell S.H., 1948. The southern tomato exchange program. Proc. Am. Soc. Hort. Sci. 52: 375-382.

30

31

Figure 3: Pedigree of ‘MR4’ & ‘NC 72 TR 4-4’ (and of ‘Venus’ and ‘Saturn’) (North Carolina State University material) (Taken from Daunay, 1977, based on W.R. Henderson & E. Echandi, pers. com. to H. Laterrot in the 1970s, and on Hendersons & Jenkins, 1972)

32

33

34

35

36

37

Figure 9 : Pedigree of ‘CRA74’, ‘Carette’ & ‘Caraibo’, (INRA, Guadeloupe material) (Taken from Anais, 1986)

38

39

40

41

Research Papers TGC REPORT VOLUME 60, 2010 Preliminary Observations on the Effectiveness of five Introgressions for Resistance to Begomoviruses in Tomatoes Luis Mejía, Rudy E. Teni, Brenda E. García, Ana Cristina Fulladolsa, and Luis Méndez, Facultad de Agronomía, Universidad de San Carlos de Guatemala; Sergio Melgar, Escuela de Biología, Universidad de San Carlos de Guatemala and Douglas P. Maxwell, Department of Plant Pathology, University of Wisconsin-Madison Introduction: Tomato-infecting begomoviruses have remained a major constraint to tomato production in many parts of the tropical and sub-tropical regions. Management strategies have involved i) extensive use of insecticides to control the whitefly vector, Bemisia tabaci, ii) production of virus-free transplants, iii) host-free periods, iv) use of whitefly-proof fabric cover in micro- and macro-tunnels for 30 or more days, and v) use of moderately resistant hybrids. Over the past decade the availability of moderately resistant hybrids has increased and most private seed companies devote resources to the incorporation of begomovirus-resistance genes into their new generation of hybrids.

Several genes for resistance to begomoviruses have been identified in the last two decades (Ji et al., 2007). Zamir et al. (1994) described the first resistance gene, Ty1, originating from the wild species Solanum chilense accession LA1969, to a region located between 4 cM and 10 cM in the short arm of chromosome 6. Later, gene, Ty2, was incorporated into the genome of the tomato from the wild species Solanum habrochaites by Hanson et al. (2000) and the introgression was located between 84 and 91 cM in chromosome 11. More recently, Ji et al. (2007b) reported gene, Ty3, in an introgression derived from S. chilense accession LA2779 and located at 19 to 25 cM in chromosome 6. An introgression in this same region from S. chilense accession LA1932 was designed Ty3a (Ji et al., 2007a). The resistant lines from the accession LA1932 had yet another gene, Ty4, which was recently located in the upper half of chromosome 3 (Ji et al., 2008) near 82 cM (Maxwell, personal communication). A major QTL, Ty5, was recently mapped in the TY172 breeding line with introgressions from S. peruvianum from approximately 16 to 46 cM on chromosome 4 (Anbinder et al., 2009). The potential for pyramiding begomovirus-resistance introgressions from different accessions of S. chilense, from different wild species such as S. peruvianum or S. habrochaites was discussed by Vidavski et al. (2008). Using several sources of resistance in one hybrid may overcome the selection of virus variants with novel genome combinations that may be more aggressive on tomatoes with one begomovirus-resistance introgression (García-Andrés et al., 2009). Because of the need to understand how different resistance genes respond to begomoviruses, this project was initiated to evaluate the effectiveness of begomovirus-resistance introgressions (Ty1, Ty2, Ty3, Ty3a, and Ty4) for conferring resistance to begomoviruses in a field trial in Guatemala. It is expected that this information will

42

provide a rationale for pyramiding begomovirus-resistance introgressions into hybrids that will have more durable resistance. Materials and Methods:

Germplasm: The breeding lines used in these experiments were: Gh13, homozygous for Ty3; Gc143-2, homozygous for Ty1 and Ty3; Gc171, homozygous for Ty3a and Ty4 (provided by J. Scott, University of Florida); CLN2116 homozygous for Ty2 (provided by P. Hanson, The World Vegetable Center) and Gh188-2, also homozygous for Ty2, and Gc21-a, homozygous for Ty3a and lacking Ty4. Gh13 was selected from the hybrid, Favi9 (Vidavsky and Czosnek, 1988). Gc143-2 was selected from a cross of Gc9 (provided by J. Scott, University of Florida) by a susceptible commercial hybrid. Gc21-a was obtained by several cycles of selfing from a cross between Gc171 with a susceptible commercial hybrid and an individual F2 plant. Gh188-2 was selected from a commercial hybrid with Ty2 introgression. The susceptible germplasm, HUJ-VF, without any of the introgressions for resistance, was used in several crosses. HUV-VF was provided by F. Vidavsky, The Hebrew University of Jerusalem. The F2 population used in the segregation of genes Ty3a and Ty4 was obtained from a hybrid produced by a cross between the resistant line Gc171 and a slightly resistant inbred (Gh44).

PCR Methods for detection of the introgressions associated with resistance to begomoviruses: DNA extraction was as reported by García et al. (2008). PCR-based molecular markers have been developed for the detection of Ty2, Ty3, Ty3a, and Ty4. Garcia et al. (2007) developed a co-dominant SCAR (Sequence Characterized Amplified Region) marker from the RFLP T0302 marker at 89 cM for the detection of the Ty2 introgression. The Ty3 and Ty3a introgressions were monitored with the co-dominant SCAR marker P6-25 (Ji et al., 2007a). The Ty4 introgression was detected with PCR primers developed by Y. Ji and J. Scott (personal communication, Ji et al., 2008). The presence of the Ty1 introgression in Gc143-2 was determined by sequencing the PCR fragments associated with the RFLP TG97 marker (García and Maxwell, unpublished). Field Evaluation of disease severity for the different populations: Each entry was replicated three times with 5 plants per replication. Four-week-old seedlings were transplanted into a field near Sanarate, Guatemala, where high levels of viruliferous whiteflies were present. In this area, at least 7 bipartite tomato-infecting begomoviruses have been identified (Nakhla et al., 2004) as well as the monopartite begomovirus, Tomato yellow leaf curl virus (Mejía and Maxwell, unpublished results). Each plant was scored at about 30-days after transplanting in either January 2009 or October 2009 using a disease severity index (DSI) from zero to six: 0, no symptoms; 1, extremely slight symptoms; 2, slight symptoms; 3, moderate symptoms; 4, severe symptoms with deformed leaves; 5, severe symptoms with stunted plant; and 6, very severe symptoms, no marketable fruit and very stunted plant.

43

Development of populations with different introgressions: Combinations of introgressions Ty3a and Ty4: An F2 population was obtained

from a F1 hybrid produced from the cross between line Gc171 (Ty3a/Ty3a, Ty4/Ty4, Scott and Schuster, 2007) by a slightly resistant line (Gh44, ty3/ty3, ty4/ty4). Leaf samples were collected from 77 individual plants for molecular analysis. The genotype of these plants was determined with the PCR primers P6-25F2 and P6-25R5 (SCAR marker P6-25) for introgression Ty3a and a PCR-based marker for Ty4 introgression (Ji et al., 2008). Individual F2 plants were selected that were dominant for one or both genes (RR, Ty3a/Ty3a, Ty4/Ty4; RS, Ty3a/Ty3a, ty4/ty4; SR, ty3/ty3, Ty4/Ty4 and SS, ty3/ty3, ty4/ty4). These plants were selfed to produce F3 seeds. Twelve F3 families, Gc171, Gh44 and a susceptible commercial control C-SS (ty3/ty3, ty4/ty4) were planted in a field to determine their phenotype, i.e., the level of resistance to begomoviruses was determined in January 2009. Each entry was replicated three times with 5 plants per replication.

Combination of introgressions for Ty2 and Ty3: F2 seed was obtained from the cross between resistant line Gh13 (Ty3/Ty3, ty2/ty2, RS-C, Martin et al., 2007), and line CLN2116 (ty3/ty3,Ty2/Ty2, SR-C). The genotype of the F2 plants was determined using molecular markers and individuals with one, both or none of the introgressions for resistance (genotypes RR, Ty3/Ty3,Ty2/Ty2; SR, ty3/ty3, Ty2/Ty2; RS, Ty3/Ty3, ty2/ty2 and SS, ty3/ty3, ty2/ty2) were allowed to self. The genotype of each F3 family was verified and seventeen F3 families were transplanted into a field along with the two parental lines and a susceptible control. Each entry was replicated three times with 5 plants per replication to determine their phenotype and DSIs were taken in October 2009.

Combination of introgressions Ty1, Ty3 and Ty2: F2 seed was obtained from the cross between resistant line Gc143-2 (Ty1/Ty1-Ty3/Ty3, ty2/ty2) and line Gh188-2, (ty1/ty1-ty3/ty3, Ty2/Ty2). The genotype of the F2 plants was determined using molecular markers for Ty3 and Ty2 and individuals with one, both or none of the introgressions were identified (RR, Ty3/Ty3, Ty2/Ty2; RS, Ty3/Ty3, ty2/ty2; SR, ty3/ty3, Ty2/Ty2 and SS, ty3/ty3, ty2/ty2). The Ty1 introgression was not determined as it is linked to Ty3. F3 seed was collected from individual F2 plants of different genotypes and the F3 families were transplanted in the field. Sixteen F3 families and both parental lines were planted and each entry was replicated three times and 5 plants per replication. Plants were scored for their DSIs in October 2009.

Combination of introgressions Ty3a and Ty2: Gc21-a (RS, Ty3a/Ty3a, ty2/ty2), was crossed with line Gh188-2 (SR, ty3/ty3, Ty2/Ty2). Heterozygous individuals were evaluated for resistance (HH, Ty3a/ty3, Ty2/ty2), along with the parental genotypes. Gc21-a (RS) and Gh188-2 (SR) were also crossed to the susceptible line HUJ-VF (ty2/ty2, ty3/ty3). The genotypes of the F1 populations were verified and their resistance phenotype determined in the field in October 2009. Three hybrids [Gc21-a X Gh188-2 (HH, Ty3a/ty3, Ty2/ty2), Gc21-a X HUJ-VF (HS, Ty3a/ty3, ty2/ty2) and Gh188-2 X HUJ-VF (SH, ty3/ty3, Ty2/ty2)], two parental lines and a susceptible commercial control (SS) were planted with three replications of 5 plants each. Plants were scored for their DSIs in October 2009.

Combination of introgressions Ty3a and Ty3: Line Gc21-a (Ty3a/Ty3a) was crossed with line Gh13 (Ty3/Ty3). The level of resistance in the hybrid (Ty3a/Ty3)

44

was evaluated in relation to the parental genotypes. The genotype of the F1 families was determined and subsequently transplanted in the field for the evaluation of their phenotype of resistance. The hybrid (Gc21-a X Gh13), both parental lines, and a susceptible control were planted in with three replications of five plants each. Plants were scored for their DSIs in October 2009. Results and Discussion: Natural inoculation with begomoviruses in a field in Guatemala with high populations of the whitefly vector was used to evaluate various combinations of begomovirus-resistance introgressions for their effectiveness to provide resistance to multiple begomoviruses. Evaluations of the introgressions Ty3a and Ty4: The highly resistant inbred line, Gc171 (Ty3a/Ty3a, Ty4/Ty4), was crossed with a slightly resistant genotype (Gh44, ty3/ty3, ty4/ty4) (Garcia, et al., 2008a). Among the 77 F2 plants analyzed, 17 were found to be homozygous Ty3a/Ty3a, 12 homozygous ty3a/ty3a, and 48 heterozygous Ty3a/ty3a. With relation to Ty4 introgression, 10 plants were found to be homozygous Ty4/Ty4, 42 homozygous ty4/ty4, and 25 heterozygous Ty4/ty4. Seven F3 families were evaluated: three with genotype ty3/ty3, Ty4/Ty4 (SR), and four with genotype Ty3a/Ty3a, ty4/ty4 (RS) (Fig. 1).

Fig. 1. Disease severity index (DSI) and standard errors for F3 families of genotypes RR (4 families Ty3a/Ty3a, Ty4/Ty4), RS (4 families, Ty3a/Ty3a, ty4/ty4), SR (4 families, ty3/ty3, Ty4/Ty4), C-SS (commercial susceptible control), R-RR (parent, Gc171), S-SS (Gh44).

0

1

2

3

4

5

6

7

RR RS SR R-RR S-SS C-SS

DSI

F3 Families and Parents

45

The average DSI for the RS F3 families with the Ty3a introgression was 2.1, while the SR F3 families with the Ty4 introgression was 3.6, which is similar to the slightly resistant parent, Gh44 (S-SS). These results indicate that there is a larger effect on resistance by Ty3a than by Ty4 to the multiple bipartite begomoviruses present in this area. This is consistent with the report by Ji et al. (2008) where the Ty4 introgression had a lesser effect on resistance to Tomato yellow leaf curl virus than the Ty3a introgression.

Combinations of introgression Ty2 and Ty3: Seventeen F3 families were obtained from individual F2 plants arising from the cross of Gh13 X CLN2116 (F1, Ty3/ty3, Ty2/ty2). The F3 families were evaluated for symptom development 36 days after transplanting (Fig. 2).

Fig. 2. Disease severity index (DSI) with standard errors for the F3 families of genotypes RR (6 families, Ty3/Ty3, Ty2/Ty2), SR (5 families, ty3/ty3, Ty2/Ty2), RS (3 families, Ty3/Ty3, ty2/ty2), SS (3 families, ty3/ty3, ty2/ty2), SR-C (CLN2116, ty3/ty3, Ty2/Ty2), RS-C (Gh13, Ty3/Ty3, ty2/ty2) and SS-C (susceptible commercial hybrid control, ty3/ty3, ty2/ty2).

0

1

2

3

4

5

6

RR SR RS SS SR-C RS-C SS-C

DS

I

F3 Families and Parents

46

The parents had average DSIs of 0.6 and 3.8 for Gh13 and CLN2116, respectively. A commercial hybrid with neither introgression had a DSI of 5.2. The average DSI of the 6 RR families was 1.6, which was greater than the most resistant parent, Gh13 (DSI 0.6), and was similar to the DSI for the 3 RS families, which had an average DSI of 1.4. The average DSI of the 5 SR families (DSI 2.9) was similar to that for the 3 SS families (DSI 2.6).

These results indicate that Ty3 introgression from Gh13 was mainly responsible for the observed level of virus resistance in the F3 families and adding the Ty2 introgression resulted in no increase in virus resistance. The susceptible control had a DSI of 5.1, while the 3 F3 families without the introgressions (ty3/ty3, ty2/ty2) had a DSI of 2.6, indicating the presence of other unknown genes for resistance. Of interest was the range of average DSIs within families of one genotype (Fig. 3). For the 6 RR families, the average DSI ranged from 0.8 to 2.3. For the 3 SS families the range was also great, 1.8 to 3.6, for the 3 RS the range was 1.1 to 1.9. For the 5 SR families the range was 2.1 to 3.2. One explanation for these differences among F3 families with the same genotype for the two introgressions could be that there was segregation of other modifying resistance genes in the F2 plants used to generate the F3 families.

Fig. 3. Disease severity index (DSI) and standard errors for the individual F3 families with the genotypes as listed in Fig. 2 along with the parents and susceptible commercial control.

0

1

2

3

4

5

6

DS

I

F3 Families and Parents

47

Combination of introgressions Ty1-Ty3 and Ty2: Individual F2 plants of different genotype obtained from the F1 (Ty1-Ty3/ty1-ty3, Ty2/ty2) of the cross of Gc143-2 X Gh188-2 were selfed to produce 16 F3 families. The DSI for these families was determined in the field (Fig. 4).

Fig. 4. Average disease severity index (DSI) and standard errors for 16 F3 families of genotypes RR (5 families, Ty1-Ty3/Ty1-Ty3, Ty2/Ty2), RS (3 families, Ty1-Ty3/Ty1-Ty3, ty2/ty2), SR (3 families, ty1-ty3/ty1-ty3, Ty2/Ty2), SS (5 families, ty1-ty3/ty1-ty3, ty2/ty2), and the parents: RS-C (Gc143-2, Ty1-Ty3/Ty1-Ty3, ty2/ty2) and SR-C (Gh188-2, ty1-ty3/ty1-ty3, Ty2/Ty2).

0

1

2

3

4

5

6

RR RS SR RS-C SR-C SS

DS

I

F3 Families and Parents

48

The average DSI for RS families with the T1-Ty3 introgression was 1.7 while DSI for those SR families with the Ty2 introgression was 4.8. RR families with both introgressions for resistance had an average DSI of 1.4 and those with neither introgression had DSI of 3.6. There was no difference in the average DSI for the RR and SR families. Surprisingly, the DSI for the SR families was higher than the DSI for the SS families. These observations indicated that Ty2 introgression provides no effective resistance to these begomoviruses.

An important observation was that there was a considerable range in the average DSI for families with the same genotype for these two introgressions (Fig. 5). For example, one RR family had a DSI of 3.2, which was greater than the other four RR families (range of 0.3 to 1.4). Within the RS families, one family had a DSI of 0.2 and another RS family had a DSI of 3.8. The genotype for the markers in these families, i.e., the presence or absence of the PCR fragments corresponding to the S. lycopersicum size or the introgression size, was reconfirmed by additional testing in the laboratory. Explanations to consider are that there could have been a recombination event between the PCR marker and the resistance gene for these introgressions or that other genes controlling resistance were segregating within the F2 plants.

Fig. 5. Average disease severity index (DSI) and standard errors for individual F3 families for each genotype and the two parents. See Fig. 3 for codes.

-1

0

1

2

3

4

5

6

DS

I

F3 Families and Parents

49

Combinations of genes Ty3a and Ty2: Line Gc21-a (DSI 0.7), homozygous for Ty3a, was crossed to line Gh188-2 (DSI 5.2), homozygous for Ty2; and the DSI for the F1 (HH, Ty3a/ty3, Ty2/ty2) was 0.4. Lines Gc21-a and Gh188-2 were also crossed with susceptible line HUJ-VF (ty3/ty3, ty2/ty2). The resulting heterozygous F1‟s, Ty3a/ty3, ty2/ty2 (HS) and ty3/ty3, Ty2/ty2 (SH) had DSIs of 2.9 and 4.2, respectively (Fig. 6). Thus, the Ty2 gene along either in a homozygous or heterozygous condition did not confer an adequate level of resistance. The Ty3a introgression in the heterozygous condition conferred a moderate level of resistance, but when both resistance introgressions were present in the heterozygous condition the F1 was highly resistant.

Fig. 6. Average disease severity index (DSI) and standard errors for the heterozygous genotypes HH (Ty2/ty2, Ty3a/ty3), SH (ty3/ty3, Ty2/ty2), HS (Ty3a/ty3, ty2/ty2) and homozygous parental lines RS (Ty3a/Ty3a, ty2/ty2), SR (ty3/ty3, Ty2/Ty2), and susceptible commercial control SS (ty2/ty2, ty3/ty3).

0

1

2

3

4

5

6

HH SH HS SR RS SS

DSI

F1 families and Parents

50

Combinations of introgressions Ty3a and Ty3: Line Gc21-a (Ty3a/Ty3a, DSI 0.3) was crossed to line Gh13 (Ty3/Ty3, DSI 0.4) and DSI for the F1 was 0.2. The average DSI of the commercial susceptible hybrid was 4.0 (Fig. 7).

Fig. 7. Average disease severity index (DSI) and standard errors for heterozygous genotypes Gc21-a X Gh13 (Ty3a/Ty3), Gc21-a (Ty3a/Ty3a), Gh13 (Ty3/Ty3) and commercial susceptible hybrid, (com. hybrid).

These results indicate that the combination of introgressions Ty3 and Ty3a confers a high level of resistance (DSI 0.2), and is not different than either of the resistant parents.

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

21-a 21-a X Gh13 Gh13 Com. Hybrid

DSI

F1 families and Parents

51

Conclusions: It should be recognized that not all of the comparisons contained all of the genotypes that would have been useful for making conclusions, that viruliferous whiteflies populations would have existed at different times, that several bipartite begomoviruses as well as Tomato yellow leaf curl virus would have been present, and that weather conditions may have impacted symptom expression. In general, the following observations were considered important concerning the various introgressions: 1) Ty3a was a more effective source of resistance than Ty4. 2) Ty2 either in the homozygous or heterozygous condition was not an effective source of resistance unless it was associated with another resistance gene, such as Ty3a. 3) The heterozygous families with Ty3a were not as resistant as families that were homozygous for this introgression (Fig. 6). 4) F1 hybrids with two different introgressions were highly resistant, such as Ty3a and Ty2 (Fig. 6) or Ty3a and Ty3 (Fig. 7). 5) There was considerable variation for the DSIs among the F3 families with same genotype for an introgression, which indicated that other genes were important in conditioning resistance. This was most notable in the F3 families from the cross of Gc143-2 by Gh188-2 (Fig. 5). Others have reported that several genes are involved in highest level of begomovirus resistance expression (Anbinder et al., 2009; Vidavsky and Czosnek, 1998; Zamir et al., 1994) and Anbinder et al. (2009) found that there were minor QTLs for resistance associated with both the resistant and susceptible parent used in the cross for their molecular analysis of resistance. 6) For the near future it is not enough to only use molecular markers for breeding for begomovirus resistance, but field evaluations with high levels of viruliferous whiteflies will continue to be an important part of any tomato breeding program for begomovirus resistance. Acknowledgements: This project was funded in part by grant FODECYT 54-07 to L. Mejía, by Facultad de Agronomía, Universidad de San Carlos, and by the College of Agricultural and Life Sciences, University of Wisconsin-Madison. Appreciation is expressed to Semillas Tropicales, S.A. for assistance with production of the seedlings and the field facilities.

52

Literature Cited: Abinder, I., Reuveni, M., Azari, R., Paran,I., Nahon, S., Shlomo, H., Chen, L., Lapidot,

M., and Levin, I. 2009. Molecular dissection of Tomato leaf curl virus resistance in tomato line TY172 derived from Solanum peruvianum. Theor. Appl. Genet. 119:519-530.

Garcia, B.E., Graham, E., Jensen, K.S., Hanson, P., Mejía, L., and Maxwell, D.P. 2007. Co-dominant SCAR for detection of the begomovirus-resistance Ty2 locus derived from Solanum habrochaites in tomato germplasm. Tom. Gen. Coop. Rept. 57:21-24.

Garcia, B.E., Barillas, A.C., Maxwell, D.P., and Mejia, L. 2008a. Genetic analysis of an F2 population for the segregation of two introgressions associated with the begomovirus-resistant parent, Gc171. Tomato Genetic Cooperative Report 58:18-21.

García, B.E., Mejía, L, Melgar, S., Teni, R., Sánchez-Pérez, A., Barillas, A.C., Montes, L., Keuler, N.S., Salus, M.S., Havey, M.J., and Maxwell, D.P. 2008b. Effectiveness of the Ty-3 introgression for conferring resistance in F3 families of tomato to bipartite begomoviruses in Guatemala. Tom. Genetic Coop. 58:22-28.

García-Andrés, S., Tomás, D.M., Navas-Castillo, J., and Moriones, E. 2009. Resistance-driven selection of begomoviruses associated with the tomato yellow leaf curl disease. Virus Research 146:66-72.

Hanson, P.M., Bernacchi, D., Green, S., Tanksley, S.D., Muniyappa, V., Padmaja, A.S., Chen, H.M., Kuo, G., Fang, D., and Chen, J.T. 2000. Mapping of a wild tomato introgression associated with tomato yellow leaf curl virus resistance in a cultivated tomato line. J. Amer. Soc. Hort. Sci. 125:15-20.

Ji, Y., Salus, M.S., van Betteray, B., Smeets, J., Jensen, K., Martin, C.T., Mejía, L., Scott, J.W., Havey, M.J., and Maxwell, D.P. 2007a. Co-dominant SCAR markers for detection of the Ty-3 and Ty-3a loci from Solanum chilense at 25 cM of chromosome 6 of tomato. Rept. Tomato Genetic Coop. 57:25-28.

Ji, Y., Schuster, D.J., and Scott, J.W. 2007b. Ty3, a begomovirus resistance locus near the Tomato yellow leaf curl virus resistance locus Ty-1 on chromosome 6 of tomato. Mol. Breeding 20:271-284.

Ji, Y., Scott, J.W., Hanson, P., Graham, E., and Maxwell, D.P. 2007c. Sources of resistance, inheritance and location of genetic loci conferring resistance to members of the tomato-infecting begomoviruses. In: Czosnek, H. (ed), Tomato yellow leaf curl virus Disease. Springer, The Netherlands, pp. 343-362.

Ji, Y., Scott, J.W., Maxwell, D.P., and Schuster, D.J. 2008. Ty-4, a Tomato yellow leaf curl virus resistance gene on chromosome 3 of tomato. Tomato Genet. Coop. Rep. 58:29-31.

Martin, C.T., Salus, M.S., Garcia, B.E., Jensen, K.S., Montes, L., Zea, C., Melgar, S., El Mehrach, K., Ortiz, J., Sanchez, A., Havey, M.J., Mejía, L., and Maxwell, D.P. 2007. Evaluation of PCR-based markers for scanning tomato chromosomes for introgressions from wild species. Rept. Tomato Genetic Coop. 57:31-34.

Mejía, L., Teni, R.E., Vidavski, F., Czosnek, H., Lapidot, M., Nakhla, M.K., and Maxwell, D.P. 2005. Evaluation of tomato germplasm and selection of breeding lines for resistance to begomoviruses in Guatemala. Acta Hort. 695:251-255.

53

Nakhla, M.K., Sorenson, A., Mejía, L., Ramírez, P., Karkashian, J.P., and Maxwell, D.P. 2005. Molecular Characterization of Tomato-Infecting Begomoviruses in Central America and Development of DNA-Based Detection Methods. Acta Hort. 695:277-288.

Scott, J.W., and Schuster, D.J. 2007. Gc9, Gc171, and Gc173 begomovirus resistant inbreds. Tomato Cooperative Genetics Report 57:45-46.

Vidavski, F., Czosnek, H., Gazit, S., Levy, D., and Lapidot, M. 2008. Pyramiding of genes conferring resistance to Tomato yellow leaf curl virus from different wild tomato species. Plant Breeding 127:625-631.

Vidavsky, F., and Czosnek, H. 1998. Tomato breeding lines immune and tolerant to tomato yellow leaf curl virus (TYLCV) issued from Lycopersicon hirsutum. Phytopathology 88:910-914.

Zamir, D., Michelson, I., Zakay, Y., Navot, N., Zeidan, N., Sarfatti, M., Eshed, Y., Harel, E., Pleban, T., van-Oss, H., Kedar, N., Rabinowitch, H.D., and Czosnek, H. 1994. Mapping and introgression of a tomato yellow leaf curl virus tolerance gene, Ty-1. Theor. Appl. Genet. 88:141-146.

54

Research Papers TGC REPORT VOLUME 60, 2010 Preliminary report on association of ‘Candidatus Liberibacter solanacearum’ with field grown tomatoes in Guatemala

Luis Mejía, Amilcar Sánchez, and Luis Méndez, Facultad de Agronomía, Universidad de San Carlos de Guatemala; D. P. Maxwell, Department of Plant Pathology, University of Wisconsin-Madison; R. L. Gilberston, Department of Plant Pathology, University of California-Davis; V.V. Rivera and G.A. Secor, Department of Plant Pathology, North Dakota State University, Fargo. Introduction A new disease of tomatoes has received considerable attention in the local newspapers in Guatemala. Locally, it is referred to “Paratrioza disease”, which refers to the insect associated with symptomatic plants. The symptoms on tomatoes are flower abortion, purple margins of youngest leaves, upward cupping of leaves, thickened stems and retarded internode growth, and stunting of the plants (Fig. 1). In Mexico, a disease of tomato with similar symptoms is called permanent damage disease or permanent yellowing disease (daňo permanente del tomate; Páramo Menchaca, 2007) and Munyaneza et al. (2009) reported that Candidatus Liberibacter solanacearum was associated with these plants. This unculturable bacterium is transmitted by the tomato/potato psyllid (Bactericerca (Paratrioza) cockerelli).

Fig. 1. Typical symptoms associated with “Paratrioza disease” of tomato in Guatemala. Image taken December 2009 and shows flower abortion, purple leaf margins, cupping of leaves and thickened petioles and stems.

In a tomato field (about 0.7 ha, at 1,500 m, Department of Sacatepéquez) where symptom incidence was over 90%, samples of young leaves with typical symptoms were collected in December 2009. DNA was extracted at San Carlos University, Guatemala City (Garcia et al., 2007). PCR was performed at North Dakota State

55

University with 16s rRNA primers,CLi.po.F/O12c and PCR fragments sequenced using protocols reported by Secor et al. (2009). A PCR fragment of about 1,100 bp was obtained and sequenced in both directions. After correction by visual proofreading, a 966-nt region (contact D.P. Maxwell for sequence) was submitted to a BLAST analysis at the National Center for Biotechnology Information data base; and a 100% nucleotide identity was obtained with the 16s-23s rRNA intergenic spacer region for „Candidatus Liberibacter solanacearum’ from potato from Guatemala (FJ395205, Secor et al., 2009) and tomato from Sinaloa, Mexico (FJ957897, Munyaneza et al., 2009b). These sequences also had a 100% nucleotide identity with „Ca. L. solanacearum‟ from tomato in New Zealand (EU834130, Liefting et al., 2009a, 2009b), from bell pepper collected in Sinaloa, Mexico (FJ957896, Munyaneza et al., 2009a) and „Candidatus Liberibacter psyllaurous‟ (synonym „Ca. L. solanacearum‟, see discussion in Secor et al., 2009) from Zebra chip symptomatic potatoes in California (FJ498802, Crosslin and Bester, 2009). In Central America, Rehman et al. (2010) report the widespread occurrence of „Ca. L. solanacerum‟ (GQ926922) and its potato psyllid vector in potato fields in Honduras. Pair wise comparison of this sequence from „Ca. L. solanacearum‟ from potato with the Guatemalan „Ca. L. solanacearum’ sequence from tomato showed that there was a difference of two SNP between the sequences (678 nt), and this might indicate a different geographic origin of these two pathogens. Subsequently to samples collected in December 2009, samples were collected from tomatoes with either typical begomovirus symptoms and/or purple/yellowing symptoms on younger leaves in the Department of El Progresso in March 2010. The samples were prepared for transport to the University of California-Davis using AgDia absorption strips. This is a method to capture total nucleic acids from plant tissue in which sap is prepared from the target tissue, applied to an absorbent matrix on the end of a plastic 'stick', and allowed to dry prior to transport. DNA extracts were prepared from the 12 samples and tested for the presence of Liberobacter and begomovirus infection by PCR. 'Ca. Liberibacter sp.' was detected in 4 of 12 samples, and these samples showed symptoms of stunted and distorted growth; older leaves were yellow and brittle and younger leaves were upcurled with yellowing and vein purpling. Begomovirus infection was detected in all 12 samples. Additionally, 49 tomato samples were collected from March to June, 2010 and assayed for Liberibacter sp. using PCR primers CLi.po.F/O12c (Secor et al., 2009). The expected size fragment for Liberibacter sp. was obtained from 17 samples and no fragments were obtained with the other 32 samples. Positive PCR samples were collected from the following Departments: Sacatepéquez, Guatemala and Baja Verapaz,. These PCR fragments will be sequenced for more definitive identification. Universal PCR primers for phytoplasma (Smart et al., 1996) were used with the DNA samples from tomato collected in Guatemala and all were negative. Because of the purple top symptoms, phytoplasma were originally suspected as being present.

In the last two years, there has been considerable effort devoted to understanding the etiology of Zebra Chip (see Secor et al., 2009) and the new Candidatus Liberibacter sp. associated with solanaceous plants (see Liefting et al., 2009a, 2009b). These efforts plus the report here indicate that this unculturable bacterium transmitted by the tomato/potato psyllid will result in a serious disease for

56

tomatoes and peppers grown in the field or greenhouse (Brown et al., 2010) in Central America.

No universally accepted name exists for this disease on tomatoes, as illustrated by the use of different names: tomato vein-greening in Arizona (Brown et al., 2010), permanent yellowing in Mexico (Munyaneza et al., 2009b) and Paratrioza disease in Guatemala. At least from our observations in Guatemala several symptoms are notable: flower abortion, purpling of the leaf margins, stem thickening, yellowing of younger leaves, and stunting. An internet search reveals that there is a substantial number of sources using „permanente del tomate‟ or permanent yellowing disease of tomato. Thus, it is proposed that this diseased be named Liberibacter permanente del tomate or Liberibacter yellowing disease of tomato. This would distinguish this disease from psyllid yellows (Brown et al., 2010).

57

Literature Cited: Brown, J.K., Rehman, M., Rogan, D., Martin, R.R., and Idris, A.M. 2010. First report of

„Candidatus Liberibacter psyllaurous‟ (synonym „Ca. L. solanacearum‟ associated with „tomato vein-greening‟ and „tomato psyllid yellows‟ dieseses in commercial greenhouses in Arizona. Plant Dis. 94:376.

Crosslin, J.M., and Bester, G. 2009. First report of „Candidatus Liberibacter psyllasurous‟ in Zebra chip symptomatic potatoes from California. Plant Dis. 93:551.

Garcia, B.E., Graham, E., Jensen, K.S., Hanson, P., Mejía, L., and Maxwell, D.P. 2007. Co-dominant SCAR for detection of the begomovirus-resistance Ty2 locus derived from Solanum habrochaites in tomato germplasm. Tom. Gen. Coop. Rept. 57:21-24.

Liefting, L.W., Weir, B.S., Pennycook, S.R., and Clover, G.R.G. 2009a. „Candidatus Liberibacter solanacearum‟ associated with plants in the family Solanaceae. Internat. J. System. and Evolut. Microbiol. 59:2274-2276.

Liefting, L.W., Sutherland, P.W., Ward, L.I., Paice, K.L., Weir, B.S., and Clover, G.R.G. 2009b. A new „Candidatus Liberibacter‟ species associated with diseases of Solanaceous crops. Plant Dis. 93:208-214.

Munyaneza, J.E., Sengoda, V.G., Crosslin, J.M., Garzón-Tiznado, J.A., and Cardenas-Valenzuela, O.G. 2009a. First report of „Candidatus Liberibacter solanacearum‟ in pepper plants in México. Plant Dis. 93:1079.

Munyaneza, J.E., Sengoda, V.G., Crosslin, J.M., Garzón-Tiznado, J.A., and Cardenas-Valenzuela, O.G. 2009b. First report of „Candidatus Liberibacter solanacearum‟ in tomato plants in México. Plant Dis. 93:1079.

Páramo Menchaca, V. Estrategia integrada: control de Paratrioza, pulgón saltador o psílido de la papa y el tomate. Productores de Hortalizas. April, 2007.

Rehman, M., Melgar, J.C., Rivera, J.M., Idris, A.M., and Brown, J.K. 2010. First report of „Candidatus Liberibacter psyllaurous‟ or „Ca. Liberibacter solanacearum‟ associated with severe foliar chlorosis, curling, and necrosis and tuber discoloration of potato plants in Honduras. Plant Dis. 94:376.

Secor, G.A., Rivera, V.V., Abad, J.A., Clover, G.R.G, Liefting, L.W., Li. X., and De Boer, S.H. 2009. Association of „Candidatus Liberibacter solanacearum‟ with Zebra Chip disease of potato established by graft and psyllid transmission, electron microscopy, and PCR. Plant Dis. 93:574-583.

Smart, C.D., Schneider, B., Blomquist, C.L., Guerra, L.J., Harrison, N.A., Ahrens, U., Lorenz, K.-H., Seemuller, E., and Kirkpatrick, B.C. 1996. Phytoplasm-specific PCR primers based on sequences of the 16s-23s rRNA spacer region. Applied and Environ. Microbiol. 62:2988-2993.

58

Research Papers TGC REPORT VOLUME 60, 2010 Study of epidermal cell size of petals and stamens in tomato species and hybrids using confocal laser-scanning microscopy Christopher Lofty, Julian Smith, Pravda Stoeva-Popova Department of Biology, Winthrop University, Rock Hill SC 29733 E-mail: stoevap@winthrop.edu Introduction

The phenomenon of cytoplasmic male sterility (CMS) has been described and the genetics underlying the phenomemon studied in many species. Whether arising spontaneously, as the result of mutations, or through alloplasmic incompatibilities in interspecific crosses, the main effect of CMS is on the development of stamens and pollen, leading to aberrant stamens with no pollen or aborted pollen (Kaul 1988). Other changes correlated with the CMS phenotype are changes in the second whorl affecting petal size and color (Andersen 1963, 1964; Petrova et al. 1999; Farbos et al. 2001; Leino et al. 2003)

In the tomato, CMS does not occur naturally. CMS has been reported in interspecies hybrids. Andersen (1963, 1964) reported the emergence and increase of pollen abortion in F1 and backcrosses of the crosses between Solanum lycopersicum, S. cheesmaniiae (formerly L. chesmanii f. typicum and f. minor) or S. habrochaites (formerly L. hirsutum f. glabratum) used as pistillate parents and S. pennellii as the recurrent pollinating parent. Pleiotropic effects of the CMS phenotype included the reduction of anther length and size, and the lengthening of the filaments. The anther size was negatively correlated to the percent of aborted pollen.

Similar results were observed by Valkova-Atchkova (1980) in crosses involving S. peruvianum as pistillate parent and S. pennellii and S. habrochaites (formerly L. hirsutum f. typicum) as pollinating parents. Further introgression of the nuclear genome of the recurrent parents confirmed the stability of the CMS phenotype over many generations (Petrova et al. 1999, Stoeva et al. 2007).

As a preliminary step to dissecting morphological (and underlying genetic) changes occurring in the CMS phenotype, this study has focused on the comparative analysis of the size of epidermal cells from abaxial and adaxial sides of petals and stamens of mature flowers from CMS-pennellii line (0% stainable pollen), the isonuclear S. pennellii (100% pollen fertility), and the cultivated tomato S. lycopersicum. Materials and methods Plant material

In the study the following genotypes were used: Solanum lycopersicum (cv. Merkurii), Solanum pennellii (LA716), and CMS-pennellii (CMS line) previously described in Petrova et al. (1998) and Radkova (2002). To determine the size of the epidermal cells, fully expanded flowers in stage 20 according to the classification of Burkhin et al. (2003) were collected from plants grown in the same environmental

59

chamber with 24C/ 20C day/night temperature and 18/6 hours day/night photoperiod (Fig. 1). Tomato flower preparation for confocal microscopy: A 0.1% aqueous solution of calcofluor white (SIGMA) was made (Pringle 1991). Excess solution was kept in the freezer and defrosted as needed. Excised floral structures (petals, anthers or filaments) were immersed in approximately 5 ml of 0.1% calcofluor solution and placed under vacuum until rapid boil was achieved. The sunken specimens were allowed to soak for 20 minutes in the dark and then were rinsed in dH2O for 10 minutes. Specimens were dissected and placed on slides. For the anthers, spacers of approximately 0.5 mm – 1 mm thick were used to ensure the specimens were not smashed by the coverslip. Coverslips were placed on top and fixed with nail polish at each corner. Calcofluor staining of epidermal cell walls was observed using the DAPI channel of the confocal laser-scanning microscope (CLSM) (FV1000). Images were taken at the planes in which cell wall margins contacted each other. Cells were

counted in image frames measuring from 150 m X 150 m to 300 m X 270 m, with the frame size adjusted as needed to include from five to twenty cells in each frame.

Ten to fifteen cells from both sides (abaxial and adaxial) of each floral structure

were measured using ImageJ (Rasband, 2009) to obtain cell areas (in m2). The ANOVA and Tukey's LSD statistical tests (Minitab, Inc.) were performed separately for each epidermal floral surface across genotypes to determine if any significant difference existed between the mean surface areas of studied epidermal cells. Results Petal abaxial cell surface area was significantly different among the three genotypes (p < 0.001) with CMS line having the largest mean cell size and S. pennellii having the smallest (Figs. 2, 3). Petal adaxial cell sizes differed significantly with S. lycopersicum and S. pennelli both being larger than the CMS line (Fig 3). Although no statistical test was done, abaxial cells were smaller than the adaxial in the two species S. pennellii and S. lycopersicum. On the contrary, the abaxial cells in the CMS line were larger in comparison to the adaxial cells.

Epidermal cell area on both the adaxial and the abaxial surfaces of the anthers was significantly different (p< 0.001) among the three genotypes (Figs.4, 5), with S. lycopersicum having the largest mean cell size for both, and the CMS line, the smallest.

Among the filament epidermal cells, the abaxial cells in the CMS line and cultivated tomato were not significantly different, but both were significantly bigger than those in S. pennellii (p< 0.001) (Fig.6, 7). There were no significant differences in the mean size of the epidermal cells on the adaxial surface of the filaments of the three strains (p=0.252) (Fig.5, 6). There was a large variation (Fig. 7) in the size of the adaxial epidermal cells in S. pennellii and the CMS line, while the size of the adaxial cells of filament in the cultivated tomato showed less variation. Discussion

According to published data, cytoplasmic male sterility has a considerable effect on the size and color of petals and stamens (Kaul 1988, Farbos et al. 2001, Leino et al. 2003). Our previous studies had also shown that in comparison to S. pennellii, the CMS

60

flowers have smaller and lighter green petals and anthers and longer filaments (Fig 1; Petrova et al 1998). In addition to tomato nuclear male sterility mutants (reviewed in Gorman and McCormick 1997) ), similar changes of flower structures have been reported for tomato plants grown at low temperatures (Lozano et al. 1998) and in loss-of-function transgenic plants for the B-class genes required for petal and stamen identity (De Martino et al. 2006).

The size of plant organs is determined by cell division and cell expansion and elongation. Our investigation of the size of epidermal cells shows that the smaller size of the anthers in the CMS line (Fig. 1; Stoeva-Popova et al., unpublished) can be explained largely by the reduction in the size of the cells. The epidermal cells on both surfaces of the CMS anther were significantly smaller in comparison to S. pennellii: respectively 51% smaller on the adaxial side, and 42.3% on the abaxial surface. On the other hand, no corresponding effect of cytoplasmic male sterility on cell size in the petals was observed On the abaxial surface of the petals of CMS line, the cells were 81.5% larger than in S. pennellii, while on the adaxial side, they were 32.2 % smaller. According to our measurements (Stoeva et al., unpublished data) the CMS-pennellii filaments are several magnitudes longer than the filaments of S. pennellii (Fig. 1). This is not merely a consequence of increased cells size, as the filament epidermal cells of CMS are statistically larger than those of S. pennellii only on the abaxial side. CMS-pennellii and S. pennellii share the same nuclear genome and cytoplasmic male sterility affects expression of nuclear genes ( reviewed in Linke, Börner 2005, Chase 2006). Our results above show that cytoplasmic male sterility does not equally affect the development of petals, stamens and filaments.

Our study has shown that there are significant differences in the size of epidermal cells between the two species: the red-fruited cultivated tomato and the green-fruited S. pennellii. Significant differences were determined for the epidermal cells of the anthers and petals. The epidermal cells of the anthers of the cultivated tomato were larger, which is an indication that the anthers of S. pennellii have greater number of epidermal cells. The same conclusion could be drawn from the study of the abaxial epidermal cells of the petals and the filaments. Since the two species are not closely related it will be interesting to investigate other representatives of the tomato clade and to determine if the size of epidermal cells of flower structures can be indicative of species relatedness.

Although no statistical analysis was carried out, our data show one consistent feature across all genotypes: the largest epidermal cells were observed on anthers, while the ones with the smallest surface area were epidermal cells of petals.

61

References

Andersen W.R. (1964). Evidence for plasmon differentiation in Lycopersicon. Report Tomato Genet. Coop. 14:4-6

Andersen, W.R. (1963). Cytoplasmic sterility in hybrids of Lycopersicon esculentum and Solanum pennellii. Report Tomato Genet. Coop. 13:7-8

Burkhin V., Hernould M., Gonzalez N., Chevalier C., Mouras A. (2003). Flower development schedule in tomato Lycopersicon esculentum cv. sweet cherry. Sex. Plant. Reprod. 15:311-320

Chase Ch. (2006). Cytoplasmic male sterility: a window to the world of plant mitochondrial-nuclear interactions. Trends in Genetics 23 (3):81-90

de Martino G., Pan I., Emmanuel E., Levy A., Irish V.F. (2006). Functional analysis of two tomato APETALA3 genes demonstrate diversification in their roles in regulating floral development. Plant Cell 18:1833-1845

Farbos I., Mouras A., Bereterbide A., Glimelius K. (2001). Defective cell proliferation in the floral meristems of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility. Plant Journal 26:131-142.

Gorman S.W., McCormick S. (1997). Male sterility in tomato. Critical Reviews in Plant Sciences 16(1):31-53

Kaul M.L.H (1988). Male sterility in higher plants. In: Monographs on Theor. Appl. Genet. 10. Springer Verlag Berlin

Leino M., Teixeira R., Landgren M., Glimelius K. (2003). Brassica napus lines with rearranged Arabidopsis mitochondria display CMS and a range of developmental aberrations. Theor. Appl. Genet. 106:1156-1163

Linke B., Börner T. (2005). Mitochondrial effects on flower and pollen development. Mitochondrion 5:389-402

Lozano R., Angosto T., Gomez P., payan C., Huijer P., Salinas J., Martinez-Zapater J.M. (1998). Tomato flower abnormalities induced by low temperatures are associated with changes of expression of MADS-box genes. Plant Physiology 117:91-100

Petrova M., Vulkova Z., Gorinova N., Izhar S., Firon N., Jacquemin J.-M., Atanassov A., Stoeva P. (1999). Characterization of cytoplasmic male sterile hybrid line between Lycopersicon peruvianum Mill. x Lycopersicon pennellii Corr. and its crosses with the cultivated tomato. Theor. Appl. Genet. 98:825-830

Pringle JR. (1991) Staining of bud scars and other cell wall chitin with calcofluor. Methods in Enzymology 4:732-5

Radkova M. (2002). Morphological, cytogenetic and molecular genetic studies of cytoplasmic male sterility in genus Lycopersicon. PhD Thesis, AgroBioInstitute, Sofia, Bulgaria

Rasband W.S., (2009) ImageJ. U. S. National Institutes of Health, Bethesda, Maryland, USA, http://rsb.info.nih.gov/ij.

62

Stoeva P., Dimaculangan2D., Radkova M., Vulkova Z. (2007). Towards cytoplasmic male sterility in cultivated tomato. Journal of Agricultural, Food and Environmental Sciences 1(1): http://www.scientificjournals.org/journals2007/articles/1058.htm

Valkova-Achkova Z. (1980). L. peruvianum a source of CMS. Rep. Tomato Genet. Coop. 30:36

Authors’ contributions: P.S-P. designed the experiment and provided the plant material. C.L. prepared all of the specimens and measured the specimens using techniques designed in collaboration with J.SIII. C.L. drafted the Methods and Results sections, and performed the statistical analysis under the supervision of JSIII. P.S-P. drafted the Introduction and Discussion. All three authors contributed to the final editing and read and approved the manuscript before submission. Fig.1: Flowers of Solanum lycopersicum (A), S. pennellii (B) and CMS-pennellii (C)

A B

C

63

Fig 2: CLSM photographs of epidermal cells from abaxial and adaxial surfaces of petals from flowers (genotypes and magnification as indicated on pictures)

Fig 3: Mean epidermal surface area of petals. Bars with the same letter are statistically equal

64

Fig. 4: CLSM photographs of epidermal cells from abaxial and adaxial surfaces of mature anthers (genotypes and magnification as indicated on pictures

Fig. 5: Mean epidermal surface area of anthers. Bars with the same letter are statistically equal

65

Fig. 6: CLSM photographs of epidermal cells from abaxial and adaxial surfaces of filaments from mature stamens (genotypes and magnification as indicated on pictures)

Fig 7: Mean epidermal surface area of filaments. Bars with the same letter are statistically equal

66

TGC REPORT VOLUME 60, 2010 Revised List of Wild Species Stocks Chetelat, R. T. C.M. Rick Tomato Genetics Resource Center, Dept. of Plant Sciences, University of California, One Shields Ave., Davis, CA 95616 The following list of 1,196 accessions of wild tomatoes and allied Solanum species is a revision of the list published in TGC vol. 57, 2007. Other types of TGRC stocks are catalogued in TGC 58 (monogenic mutants) and TGC 59 (miscellaneous stocks). Inactive accessions have been dropped and new collections added to the present list. The new material includes populations of wild or feral cherry tomato (S. lycopersicum ‘cerasiforme‟) from Mexico (LA4352, LA4353), a stock of S. pimpinellifolium containing the sun gene introgressed from S. lycopersicum, and populations of S. peruvianum from the Azapa valley of northern Chile (LA4445-LA4448). Seed samples will be provided, upon request, for research, breeding or educational purposes. Some accessions may be temporarily unavailable for distribution during seed multiplication. In general, only small quantities of seed will be provided: 25 seed per accession for the self-pollinated accessions, 50 for the outcrossers or facultative accessions, and 5-10 for the allied Solanum species. These seed samples should be sufficient for researchers to produce larger quantities of seed, if needed. Accessions are grown for seed increase in the UC-Davis greenhouses, except for cherry tomatoes and certain populations of S. pimpinellifolium, which are grown in the field. The population sizes used for seed multiplication depend on the mating system, and are designed to maintain genetic diversity within accessions (see guidelines at http://tgrc.ucdavis.edu). The following tables are ordered by species name, using the classification system of Peralta et al. (2008)11, but with the equivalent Lycopersicon names listed as well. Although only brief collection site data can be presented here, more detailed records are available from our website, including geographic coordinates, images, and donor information. An appendix table lists the accessions belonging to the core subsets for each species.

S. arcanum (L. peruvianum or L. peruvianum var. humifusum)

LA0378 Cascas Cajamarca Peru

LA0385 San Juan (Rio Jequetepeque) Cajamarca Peru

LA0389 Abra Gavilan Cajamarca Peru

LA0392 Llallan Cajamarca Peru

LA0441 Cerro Campana La Libertad Peru

LA1027 Cajamarca Peru

11

Peralta, I. E., D. M. Spooner, S. Knapp (2008) Taxonomy of wild tomatoes and their relatives

(Solanum sect. Lycopersicoides, sect. Juglandifolia, and sect. Lycopersicon; Solanaceae).

Systematic Botany Monographs 84: 1-186.

67

S. arcanum (L. peruvianum or L. peruvianum var. humifusum)

LA1031 Balsas Amazonas Peru

LA1032 Aricapampa La Libertad Peru

LA1346 Casmiche La Libertad Peru

LA1350 Chauna Cajamarca Peru

LA1351 Rupe Cajamarca Peru

LA1360 Pariacoto Ancash Peru

LA1394 Balsas - Rio Utcubamba Amazonas Peru

LA1395 Chachapoyas Amazonas Peru

LA1396 Balsas (Chachapoyas) Amazonas Peru

LA1626 Mouth of Rio Rupac Ancash Peru

LA1708 Chamaya to Jaen Cajamarca Peru

LA1984 Otuzco La Libertad Peru

LA1985 Casmiche La Libertad Peru

LA2150 Puente Muyuno (Rio Jequetepeque) Cajamarca Peru

LA2151 Morochupa (Rio Jequetepeque) Cajamarca Peru

LA2152 San Juan #1 (Rio Jequetepeque) Cajamarca Peru

LA2153 San Juan #2 (Rio Jequetepeque) Cajamarca Peru

LA2157 Tunel Chotano Cajamarca Peru

LA2163 Cochabamba to Yamaluc Cajamarca Peru

LA2164 Yamaluc Cajamarca Peru

LA2172 Cuyca Cajamarca Peru

LA2185 Pongo de Rentema Amazonas Peru

LA2326 Above Balsas Amazonas Peru

LA2327 Aguas Calientes Cajamarca Peru

LA2328 Aricapampa La Libertad Peru

LA2330 Chagual La Libertad Peru

LA2331 Agallapampa La Libertad Peru

LA2333 Casmiche La Libertad Peru

LA2334 San Juan Cajamarca Peru

LA2388 Cochabamba to Huambos (Chota) Cajamarca Peru

LA2548 La Moyuna (Magadalena) Cajamarca Peru

LA2550 El Tingo, Chorpampa (Rio Jequetepeque) Cajamarca Peru

LA2553 Balconcillo de San Marcos Cajamarca Peru

LA2555 Marical - Castilla La Libertad Peru

LA2565 Potrero de Panacocha a Llamellin Ancash Peru

LA2566 Cullachaca Ancash Peru

LA2582 San Juan (4x) Cajamarca Peru

LA2583 (4x)

LA2917 Chullchaca Ancash Peru

LA4316 Kuntur Wasi Cajamarca Peru

S. cheesmaniae (L. cheesmanii)

LA0166 Santa Cruz: Barranco, N of Punta Ayora Galapagos Islands Ecuador

68

S. cheesmaniae (L. cheesmanii)

LA0421 Cristobal: cliff East of Wreck Bay Galapagos Islands Ecuador

LA0422 San Cristobal: Wreck Bay, Puerto Baquerizo Galapagos Islands Ecuador

LA0428 Santa Cruz: Trail Bellavista to Miconia Zone Galapagos Islands Ecuador

LA0429 Santa Cruz: Crater in highlands Galapagos Islands Ecuador

LA0434 Santa Cruz: Rambech Trail Galapagos Islands Ecuador

LA0437 Isabela: Ponds North of Villamil Galapagos Islands Ecuador

LA0521 Fernandina: Inside Crater Galapagos Islands Ecuador

LA0522 Fernandina: Outer slopes Galapagos Islands Ecuador

LA0524 Isabela: Punta Essex Galapagos Islands Ecuador

LA0528B Santa Cruz: Academy Bay Galapagos Islands Ecuador

LA0529 Fernandina: Crater Galapagos Islands Ecuador

LA0531 Baltra: Barranco slope, N side Galapagos Islands Ecuador

LA0746 Isabela: Punta Essex Galapagos Islands Ecuador

LA0749 Fernandina: North side Galapagos Islands Ecuador

LA0927 Santa Cruz: Academy Bay Galapagos Islands Ecuador

LA0932 Isabela: Tagus Cove Galapagos Islands Ecuador

LA1035 Fernandina: Low elevation Galapagos Islands Ecuador

LA1036 Isabela: far north end Galapagos Islands Ecuador

LA1037 Isabela: Alcedo East slope Galapagos Islands Ecuador

LA1039 Isabela: Cape Berkeley Galapagos Islands Ecuador

LA1040 San Cristobal: Caleta Tortuga Galapagos Islands Ecuador

LA1041 Santa Cruz: El Cascajo Galapagos Islands Ecuador

LA1042 Isabela: Cerro Santo Tomas Galapagos Islands Ecuador

LA1043 Isabela: Cerro Santo Tomas Galapagos Islands Ecuador

LA1138 Isabela: E of Cerro Azul Galapagos Islands Ecuador

LA1139 Isabela: W of Cerro Azul Galapagos Islands Ecuador

LA1402 Fernandina: W of Punta Espinoza Galapagos Islands Ecuador

LA1404 Fernandina: W flank caldera Galapagos Islands Ecuador

LA1406 Fernandina: SW rim caldera Galapagos Islands Ecuador

LA1407 Fernandina: caldera, NW bench Galapagos Islands Ecuador

LA1409 Isabela: Punta Albermarle Galapagos Islands Ecuador

LA1412 San Cristobal: opposite Isla Lobos Galapagos Islands Ecuador

LA1414 Isabela: Cerro Azul Galapagos Islands Ecuador

LA1427 Fernandina: WSW rim of caldera Galapagos Islands Ecuador

LA1447 Santa Cruz: Darwin Station-Punta Nunez Galapagos Islands Ecuador

LA1448 Santa Cruz: Puerto Ayora, Pelican Bay Galapagos Islands Ecuador

LA1449 Santa Cruz: Darwin Station, Seismo Station Galapagos Islands Ecuador

LA1450 Isabela: Bahia San Pedro Galapagos Islands Ecuador

LA3124 Santa Fe: near E landing Galapagos Islands Ecuador

S. chilense (L. chilense)

LA0130 Moquegua Moquegua Peru

LA0294 Tacna Tacna Peru

69

S. chilense (L. chilense)

LA0456 Clemesi Moquegua Peru

LA0458 Tacna Tacna Peru

LA0460 Palca Tacna Peru

LA0470 Taltal Antofagasta Chile

LA1029 Moquegua Moquegua Peru

LA1030 Tarata Rd. Tacna Peru

LA1782 Quebrada de Acari Arequipa Peru

LA1917 Llauta (4x) Ayacucho Peru

LA1930 Quebrada Calapampa Arequipa Peru

LA1932 Minas de Acari Arequipa Peru

LA1938 Quebrada Salsipuedes Arequipa Peru

LA1958 Pampa de la Clemesi Moquegua Peru

LA1959 Huaico Moquegua Moquegua Peru

LA1960 Rio Osmore Moquegua Peru

LA1961 Toquepala Tacna Peru

LA1963 Rio Caplina Tacna Peru

LA1965 Causuri Tacna Peru

LA1967 Pachia, Rio Caplina Tacna Peru

LA1968 Cause Seco Tacna Peru

LA1969 Estique Pampa Tacna Peru

LA1970 Tarata Tacna Peru

LA1971 Palquilla Tacna Peru

LA1972 Rio Sama Tacna Peru

LA2404 Arica to Tignamar Tarapaca Chile

LA2405 Tignamar Tarapaca Chile

LA2406 Arica to Putre Tarapaca Chile

LA2731 Moquella Tarapaca Chile

LA2737 Yala-yala Tarapaca Chile

LA2739 Nama to Camina Tarapaca Chile

LA2746 Asentamiento-18 Tarapaca Chile

LA2747 Alta Azapa Tarapaca Chile

LA2748 Soledad Tarapaca Chile

LA2749 Punta Blanca Antofagasta Chile

LA2750 Mina La Despreciada Antofagasta Chile

LA2751 Pachica (Rio Tarapaca) Tarapaca Chile

LA2753 Laonzana Tarapaca Chile

LA2754 W of Chusmisa Tarapaca Chile

LA2755 Banos de Chusmisa Tarapaca Chile

LA2757 W of Chusmisa Tarapaca Chile

LA2759 Mamina Tarapaca Chile

LA2762 Quebradas de Mamina a Parca Tarapaca Chile

LA2764 Codpa Tarapaca Chile

LA2765 Timar Tarapaca Chile

70

S. chilense (L. chilense)

LA2767 Chitita Tarapaca Chile

LA2768 Empalme Codpa Tarapaca Chile

LA2771 Above Poconchile Tarapaca Chile

LA2773 Zapahuira Tarapaca Chile

LA2774 Socorama Tarapaca Chile

LA2778 Chapiquina Tarapaca Chile

LA2779 Cimentario Belen Tarapaca Chile

LA2780 Belen to Lupica Tarapaca Chile

LA2879 Peine Antofagasta Chile

LA2880 Quebrada Tilopozo Antofagasta Chile

LA2882 Camar Antofagasta Chile

LA2884 Ayaviri Antofagasta Chile

LA2887 Quebrada Bandurria Antofagasta Chile

LA2888 Loma Paposo Antofagasta Chile

LA2891 Taltal Antofagasta Chile

LA2930 Quebrada Taltal Antofagasta Chile

LA2931 Guatacondo Tarapaca Chile

LA2932 Quebrada Gatico, Mina Escalera Antofagasta Chile

LA2946 Guatacondo Tarapaca Chile

LA2949 Chusmisa Tarapaca Chile

LA2952 Camiña Tarapaca Chile

LA2955 Quistagama Tarapaca Chile

LA2980 Yacango Moquegua Peru

LA2981A Torata to Chilligua en route to Puno Moquegua Peru

LA3111 Tarata Tacna Peru

LA3112 Estique Pampa Tacna Peru

LA3113 Apacheta Tacna Peru

LA3114 Quilla Tacna Peru

LA3115 W of Quilla Tacna Peru

LA3153 Desvio Omate (Rio de Osmore) Moquegua Peru

LA3155 Quinistaquillas Moquegua Peru

LA3355 Cacique de Ara Tacna Peru

LA3356 W of Tacna Tacna Peru

LA3357 Irrigacion Magollo Tacna Peru

LA3358 Rio Arunta-Cono Sur Tacna Peru

LA3784 Rio Chaparra Arequipa Peru

LA3785 Terras Blancas Arequipa Peru

LA3786 Alta Chaparra Arequipa Peru

LA4106 Taltal Antofagasta Chile

LA4107 Catarata Taltal Antofagasta Chile

LA4108 Caleta Punta Grande Antofagasta Chile

LA4109 Quebrada Canas Antofagasta Chile

LA4117A San Pedro - Paso Jama Antofagasta Chile

71

S. chilense (L. chilense)

LA4117B San Pedro - Paso Jama Antofagasta Chile

LA4118 Toconao Antofagasta Chile

LA4119 Socaire Antofagasta Chile

LA4120 Cahuisa Tarapaca Chile

LA4121 Pachica - Poroma Tarapaca Chile

LA4122 Chiapa Tarapaca Chile

LA4127 Alto Umayani Tarapaca Chile

LA4129 Pachica (Rio Camarones) Tarapaca Chile

LA4132 Esquina Tarapaca Chile

LA4319 Alto Rio Lluta Tarapaca Chile

LA4321 Quebrada Cardones Tarapaca Chile

LA4324 Estacion Puquio Tarapaca Chile

LA4327 Pachica, Rio Camarones Tarapaca Chile

LA4329 Puente del Diablo, Rio Salado Antofagasta Chile

LA4330 Caspana Antofagasta Chile

LA4332 Rio Grande Antofagasta Chile

LA4334 Quebrada Sicipo Antofagasta Chile

LA4335 Quebrada Tucuraro Antofagasta Chile

LA4336 Quebrada Cascabeles Antofagasta Chile

LA4337 Quebrada Paposo Antofagasta Chile

LA4338 Quebrada Taltal, Estacion Breas Antofagasta Chile

LA4339 Quebrada Los Zanjones Antofagasta Chile

S. chmielewskii (L. chmielewskii)

LA1028 Casinchihua Apurimac Peru

LA1306 Tambo Ayacucho Peru

LA1316 Ocros Ayacucho Peru

LA1317 Hacienda Pajonal Ayacucho Peru

LA1318 Auquibamba Apurimac Peru

LA1325 Puente Cunyac Apurimac Peru

LA1327 Sorocata Apurimac Peru

LA1330 Hacienda Francisco Apurimac Peru

LA2639B Puente Cunyac Apurimac Peru

LA2663 Tujtohaiya Cusco Peru

LA2677 Huayapacha #1 Cusco Peru

LA2678 Huayapacha #2 Cusco Peru

LA2679 Huayapacha #3 Cusco Peru

LA2680 Puente Apurimac #1 Cusco Peru

LA2681 Puente Apurimac #2 Cusco Peru

LA2695 Chihuanpampa Cusco Peru

LA3642 Ankukunka Cusco Peru

LA3643 Colcha Cusco Peru

LA3644 Puente Tincoj Cusco Peru

72

S. chmielewskii (L. chmielewskii)

LA3645 Boca del Rio Velille Cusco Peru

LA3648 Huallapachaca Apurimac Peru

LA3653 Matara Apurimac Peru

LA3654 Casinchigua to Chacoche Apurimac Peru

LA3656 Chalhuani Apurimac Peru

LA3658 Occobamba Apurimac Peru

LA3661 Pampotampa Apurimac Peru

LA3662 Huancarpuquio Apurimac Peru

S. corneliomulleri (L. peruvianum or L. peruv. f. glandulosum)

LA0103 Cajamarquilla, Rio Rimac Lima Peru

LA0107 Hacienda San Isidro, Rio Canete Lima Peru

LA0364 9 Km W of Canta Lima Peru

LA0366 12 Km W of Canta Lima Peru

LA0444 Chincha #1 Ica Peru

LA0451 Arequipa Arequipa Peru

LA1133 Huachipa Lima Peru

LA1271 Horcon Lima Peru

LA1274 Pacaibamba Lima Peru

LA1281 Sisacaya Lima Peru

LA1283 Santa Cruz de Laya Lima Peru

LA1284 Espiritu Santo Lima Peru

LA1292 San Mateo Lima Peru

LA1293 Matucana Lima Peru

LA1294 Surco Lima Peru

LA1296 Tornamesa Lima Peru

LA1304 Pampano Huancavelica Peru

LA1305 Ticrapo Huancavelica Peru

LA1331 Nazca Ica Peru

LA1339 Capillucas Lima Peru

LA1373 Asia Lima Peru

LA1377 Navan Lima Peru

LA1379 Caujul Lima Peru

LA1473 Callahuanca, Santa Eulalia valley Lima Peru

LA1551 Rimac Valley, Km 71 Lima Peru

LA1552 Rimac Valley, Km 93 Lima Peru

LA1554 Huaral to Cerro de Pasco, Rio Chancay Lima Peru

LA1609 Asia - El Pinon Lima Peru

LA1646 Yaso Lima Peru

LA1647 Huadquina, Topara Ica Peru

LA1653 Uchumayo, Arequipa Arequipa Peru

LA1677 Fundo Huadquina, Topara Ica Peru

LA1694 Cacachuhuasiin, Cacra Lima Peru

73

S. corneliomulleri (L. peruvianum or L. peruv. f. glandulosum)

LA1722 Ticrapo Viejo Huancavelica Peru

LA1723 La Quinga Ica Peru

LA1744 Putinza Lima Peru

LA1910 Tambillo Huancavelica Peru

LA1937 Quebrada Torrecillas Arequipa Peru

LA1944 Rio Atico Arequipa Peru

LA1945 Caraveli Arequipa Peru

LA1973 Yura Arequipa Peru

LA2717 Chilca Lima Peru

LA2721 Putinza Lima Peru

LA2724 Huaynilla Lima Peru

LA2962 Echancay Arequipa Peru

LA2981B Torata to Chilligua en route to Puno Moquegua Peru

LA3154 Otora-Puente Jahuay Moquegua Peru

LA3156 Omate Valley Moquegua Peru

LA3219 Catarindo (Islay) Arequipa Peru

LA3637 Coayllo Lima Peru

LA3639 Ccatac Lima Peru

LA3664 Nazca grade Ica Peru

LA3666 La Yapa Ica Peru

S. galapagense (L. cheesmanii f. minor)

LA0317 Bartolome Galapagos Islands Ecuador

LA0426 Bartolome: E of landing Galapagos Islands Ecuador

LA0436 Isabela: Villamil Galapagos Islands Ecuador

LA0438 Isabela: coast at Villamil Galapagos Islands Ecuador

LA0480A Isabela: Cowley Bay Galapagos Islands Ecuador

LA0483 Fernandina: inside crater Galapagos Islands Ecuador

LA0526 Pinta: W Side Galapagos Islands Ecuador

LA0527 Bartolome: W side, Tower Bay Galapagos Islands Ecuador

LA0528 Santa Cruz: Academy Bay Galapagos Islands Ecuador

LA0530 Fernandina: crater Galapagos Islands Ecuador

LA0532 Pinzon: NW side Galapagos Islands Ecuador

LA0747 Santiago: Cape Trenton Galapagos Islands Ecuador

LA0748 Santiago: E Trenton Island Galapagos Islands Ecuador

LA0929 Isabela: Punta Flores Galapagos Islands Ecuador

LA0930 Isabela: Cabo Tortuga Galapagos Islands Ecuador

LA1044 Bartolome Galapagos Islands Ecuador

LA1136 Gardner-near-Floreana Islet Galapagos Islands Ecuador

LA1137 Rabida: N side Galapagos Islands Ecuador

LA1141 Santiago: N crater Galapagos Islands Ecuador

LA1400 Isabela: N of Punta Tortuga Galapagos Islands Ecuador

LA1401 Isabela: N of Punta Tortuga Galapagos Islands Ecuador

74

S. galapagense (L. cheesmanii f. minor)

LA1403 Fernandina: W of Punta Espinoza Galapagos Islands Ecuador

LA1408 Isabela: SW volcano, Cape Berkeley Galapagos Islands Ecuador

LA1410 Isabela: Punta Ecuador Galapagos Islands Ecuador

LA1411 Santiago: N James Bay Galapagos Islands Ecuador

LA1452 Isabela: E slope, Volcan Alcedo Galapagos Islands Ecuador

LA1508 Corona del Diablo Islet (near Floreana) Galapagos Islands Ecuador

LA1627 Isabela: Tagus Cove Galapagos Islands Ecuador

LA3909 Bartolome: tourist landing Galapagos Islands Ecuador

S. habrochaites (L. hirsutum, L. hirsutum f. glabratum)

LA0094 Canta-Yangas Lima Peru

LA0361 Canta Lima Peru

LA0386 Cajamarca Cajamarca Peru

LA0387 Santa Apolonia Cajamarca Peru

LA0407 Mirador, Guayaquil Guayas Ecuador

LA1033 Hacienda Taulis Lambayeque Peru

LA1223 Alausi Chimborazo Ecuador

LA1252 Loja Loja Ecuador

LA1253 Pueblo Nuevo - Landangue Loja Ecuador

LA1255 Loja (Pedestal district) Loja Ecuador

LA1264 Bucay Chimborazo Ecuador

LA1265 Rio Chimbo Chimborazo Ecuador

LA1266 Pallatanga Chimborazo Ecuador

LA1295 Surco Lima Peru

LA1298 Yaso Lima Peru

LA1347 Empalme Otusco La Libertad Peru

LA1352 Rupe Cajamarca Peru

LA1353 Contumaza Cajamarca Peru

LA1354 Contumaza to Cascas Cajamarca Peru

LA1361 Pariacoto Ancash Peru

LA1362 Chacchan Ancash Peru

LA1363 Alta Fortaleza Ancash Peru

LA1366 Cajacay Ancash Peru

LA1378 Navan Lima Peru

LA1391 Bagua to Olmos Cajamarca Peru

LA1392 Huaraz - Casma Road Ancash Peru

LA1393 Huaraz - Caraz Ancash Peru

LA1557 Huaral to Cerro de Pasco, Rio Chancay Lima Peru

LA1559 Desvio Huamantanga-Canta Lima Peru

LA1560 Matucana Lima Peru

LA1624 Jipijapa Manabi Ecuador

LA1625 S of Jipijapa Manabi Ecuador

LA1648 Above Yaso Lima Peru

75

S. habrochaites (L. hirsutum, L. hirsutum f. glabratum)

LA1681 Mushka Lima Peru

LA1691 Yauyos Lima Peru

LA1695 Cacachuhuasiin, Cacra Lima Peru

LA1696 Huanchuy to Cacra Lima Peru

LA1717 Sopalache Piura Peru

LA1718 Huancabamba Piura Peru

LA1721 Ticrapo Viejo Huancavelica Peru

LA1731 Rio San Juan Huancavelica Peru

LA1736 Pucutay Piura Peru

LA1737 Cashacoto Piura Peru

LA1738 Desfiladero Piura Peru

LA1739 Canchaque to Cerran Piura Peru

LA1740 Huancabamba Piura Peru

LA1741 Sondorilla Piura Peru

LA1753 Surco Lima Peru

LA1764 West of Canta Lima Peru

LA1772 West of Canta Lima Peru

LA1775 Rio Casma Ancash Peru

LA1777 Rio Casma Ancash Peru

LA1778 Rio Casma Ancash Peru

LA1779 Rio Casma Ancash Peru

LA1918 Llauta Ayacucho Peru

LA1927 Ocobamba Ayacucho Peru

LA1928 Ocana Ayacucho Peru

LA1978 Colca Ancash Peru

LA2092 Chinuko Chimborazo Ecuador

LA2098 Sabianga Loja Ecuador

LA2099 Sabiango to Zozoranga Loja Ecuador

LA2100 Sozorango Loja Ecuador

LA2101 Cariamanga Loja Ecuador

LA2103 Lansaca Loja Ecuador

LA2104 Pena Negra Loja Ecuador

LA2105 Jardin Botanico, Loja Loja Ecuador

LA2106 Yambra Loja Ecuador

LA2107 Los Lirios Loja Ecuador

LA2108 Anganumo Loja Ecuador

LA2109 Yangana #1 Loja Ecuador

LA2110 Yangana #2 Loja Ecuador

LA2114 San Juan Loja Ecuador

LA2115 Pucala Loja Ecuador

LA2116 Las Juntas Loja Ecuador

LA2119 Saraguro Loja Ecuador

LA2124 Cumbaratza Zamora-Chinchipe Ecuador

76

S. habrochaites (L. hirsutum, L. hirsutum f. glabratum)

LA2128 Zumbi Zamora-Chinchipe Ecuador

LA2144 Chanchan Chimborazo Ecuador

LA2155 Maydasbamba Cajamarca Peru

LA2156 Ingenio Montan Cajamarca Peru

LA2158 Rio Chotano Cajamarca Peru

LA2159 Atonpampa Cajamarca Peru

LA2167 Cimentario Cajamarca Cajamarca Peru

LA2171 El Molino Piura Peru

LA2174 Rio Chinchipe, San Augustin Cajamarca Peru

LA2175 Timbaruca Cajamarca Peru

LA2196 Caclic Amazonas Peru

LA2204 Balsapata Amazonas Peru

LA2314 San Francisco Amazonas Peru

LA2321 Chirico Amazonas Peru

LA2324 Leimebamba Amazonas Peru

LA2329 Aricapampa La Libertad Peru

LA2409 Miraflores Lima Peru

LA2552 Las Flores Cajamarca Peru

LA2556 Puente Moche La Libertad Peru

LA2567 Quita Ancash Peru

LA2574 Cullaspungro Ancash Peru

LA2648 Santo Domingo Piura Peru

LA2650 Ayabaca Piura Peru

LA2651 Puente Tordopa Piura Peru

LA2722 Puente Auco Lima Peru

LA2812 Lambayeque Lambayeque Peru

LA2855 Mollinomuna, Celica Loja Ecuador

LA2860 Cariamanga Loja Ecuador

LA2861 Las Juntas Loja Ecuador

LA2863 Macara Loja Ecuador

LA2864 Sozorango Loja Ecuador

LA2869 Matola-La Toma Loja Ecuador

LA2975 Coltao Ancash Peru

LA2976 Huangra Ancash Peru

LA3794 Alta Fortaleza Ancash Peru

LA3796 Anca, Marca Ancash Peru

LA3854 Llaguén La Libertad Peru

LA3862 Purunuma Loja Ecuador

LA3863 Sozoranga Loja Ecuador

LA3864 Yangana Loja Ecuador

LA4137 Barrio Delta, Cajamarca Cajamarca Peru

77

S. huaylasense (L. peruvianum)

LA0110 Cajacay Ancash Peru

LA1358 Yautan Ancash Peru

LA1364 Alta Fortaleza Ancash Peru

LA1365 Caranquilloc Ancash Peru

LA1981 Vocatoma Ancash Peru

LA1982 Huallanca Ancash Peru

LA1983 Rio Manta Ancash Peru

LA2068 Chasquitambo Ancash Peru

LA2561 Huallanca Ancash Peru

LA2562 Canon del Pato Ancash Peru

LA2563 Canon del Pato Ancash Peru

LA2575 Valle de Casma Ancash Peru

LA2808 Huaylas Ancash Peru

LA2809 Huaylas Ancash Peru

S. juglandifolium

LA2120 Sabanilla Zamora-Chinchipe Ecuador

LA2134 Tinajillas Zamora-Chinchipe Ecuador

LA2788 Quebrada La Buena Antioquia Colombia

LA3322 Quito Pinchincha Ecuador

LA3323 Manuel Cornejo Astorga Pichincha Ecuador

LA3324 Sabanillas Zamora-Chinchipe Ecuador

LA3325 Cosanga Napo Ecuador

LA3326 Sicalpa Chimborazo Ecuador

S. lycopersicoides

LA1964 Chupapalca Tacna Peru

LA1966 Palca Tacna Peru

LA1990 Palca Tacna Peru

LA2385 Chupapalca to Ingenio Tacna Peru

LA2386 Chupapalca Tacna Peru

LA2387 Lago Aricota (Tarata) Tacna Peru

LA2407 Arica to Putre Tarapaca Chile

LA2408 Above Putre Tarapaca Chile

LA2730 Moquella Tarapaca Chile

LA2772 Zapahuira Tarapaca Chile

LA2776 Catarata Perquejeque Tarapaca Chile

LA2777 Putre Tarapaca Chile

LA2781 Desvio a Putre Tarapaca Chile

LA2951 Quistagama Tarapaca Chile

LA4018 Lago Aricota Tacna Peru

LA4123 Camina Tarapaca Chile

LA4126 Camina - Nama Tarapaca Chile

78

S. lycopersicoides

LA4130 Pachica (Rio Camarones) Tarapaca Chile

LA4131 Esquina Tarapaca Chile

LA4320 Rio Lluta Tarapaca Chile

LA4322 Quebrada Cardones Tarapaca Chile

LA4323 Putre Tarapaca Chile

LA4326 Cochiza, Rio Camarones Tarapaca Chile

S. lycopersicum (L. esculentum var. cerasiforme)

LA0168 New Caledonia Fr. Oceania

LA0292 Santa Cruz Galapagos Islands Ecuador

LA0349 Unknown

LA0384 Chilete (Rio Jequetepeque) Cajamarca Peru

LA0475 Sucua Morona-Santiago Ecuador

LA0476 Sucua Morona-Santiago Ecuador

LA1025 Oahu: Wahiawa Hawaii USA

LA1203 Ciudad Vieja Guatemala

LA1204 Quetzaltenango Guatemala

LA1205 Copan Honduras

LA1206 Copan Ruins Honduras

LA1207 Mexico

LA1208 Sierra Nevada Colombia

LA1209 Colombia

LA1226 Sucua Morona-Santiago Ecuador

LA1227 Sucua Morona-Santiago Ecuador

LA1228 Macas, San Jacinto de los Monos Morona-Santiago Ecuador

LA1229 Macas Plaza Morona-Santiago Ecuador

LA1230 Macas Morona-Santiago Ecuador

LA1231 Tena Napo Ecuador

LA1247 La Toma Loja Ecuador

LA1268 Chaclacayo Lima Peru

LA1286 San Martin de Pangoa Junin Peru

LA1287 Fundo Ileana #1 Junin Peru

LA1289 Fundo Ileana #3 Junin Peru

LA1290 Mazamari Junin Peru

LA1291 Satipo Granja Junin Peru

LA1307 Hotel Oasis, San Francisco Ayacucho Peru

LA1308 San Francisco Ayacucho Peru

LA1310 Hacienda Santa Rosa Ayacucho Peru

LA1311-1 Santa Rosa Puebla Ayacucho Peru

LA1311-10 Santa Rosa Puebla Ayacucho Peru

LA1311-11 Santa Rosa Puebla Ayacucho Peru

79

S. lycopersicum (L. esculentum var. cerasiforme)

LA1311-12 Santa Rosa Puebla Ayacucho Peru

LA1311-13 Santa Rosa Puebla Ayacucho Peru

LA1311-14 Santa Rosa Puebla Ayacucho Peru

LA1311-15 Santa Rosa Puebla Ayacucho Peru

LA1311-16 Santa Rosa Puebla Ayacucho Peru

LA1311-17 Santa Rosa Puebla Ayacucho Peru

LA1311-18 Santa Rosa Puebla Ayacucho Peru

LA1311-19 Santa Rosa Puebla Ayacucho Peru

LA1311-2 Santa Rosa Puebla Ayacucho Peru

LA1311-3 Santa Rosa Puebla Ayacucho Peru

LA1311-4 Santa Rosa Puebla Ayacucho Peru

LA1311-5 Santa Rosa Puebla Ayacucho Peru

LA1311-6 Santa Rosa Puebla Ayacucho Peru

LA1311-7 Santa Rosa Puebla Ayacucho Peru

LA1311-8 Santa Rosa Puebla Ayacucho Peru

LA1311-9 Santa Rosa Puebla Ayacucho Peru

LA1312-2 Paisanato Cusco Peru

LA1312-3 Paisanto Cusco Peru

LA1312-4 Paisanato Cusco Peru

LA1314 Granja Pichari Cusco Peru

LA1320 Hacienda Carmen Apurimac Peru

LA1323 Pfacchayoc Cusco Peru

LA1324 Hacienda Potrero, Quillabamba Cusco Peru

LA1328 Rio Pachachaca Apurimac Peru

LA1334 Pescaderos Arequipa Peru

LA1338 Puyo Napo Ecuador

LA1372 Santa Eulalia Lima Peru

LA1385 Quincemil Cusco Peru

LA1386 Balsas Amazonas Peru

LA1387 Quincemil Cusco Peru

LA1388 San Ramon Junin Peru

LA1420 Lago Agrio Napo Ecuador

LA1421 Santa Cecilia Napo Ecuador

LA1423 Near Santo Domingo Pichincha Ecuador

LA1425 Villa Hermosa Cauca Colombia

LA1426 Cali Cauca Colombia

LA1429 La Estancilla Manabi Ecuador

LA1453 Kauai: Poipu Hawaii USA

LA1454 Mexico

LA1455 Gral Teran Nuevo Leon Mexico

LA1456 Papantla Vera Cruz Mexico

LA1457 Tehuacan Puebla Mexico

LA1458 Huachinango Puebla Mexico

80

S. lycopersicum (L. esculentum var. cerasiforme)

LA1461 University Philippines, Los Banos Philippines

LA1464 El Progreso, Yoro Honduras

LA1465 Taladro, Comayagua Honduras

LA1467 Cali Cauca Colombia

LA1468 Fte. Casa, Cali Cauca Colombia

LA1479 Sucua Morona-Santiago Ecuador

LA1480 Sucua Morona-Santiago Ecuador

LA1481 Sucua Morona-Santiago Ecuador

LA1482 Segamat Malaysia

LA1483 Trujillo Saipan

LA1509 Tawan Sabah Borneo

LA1510 Mexico

LA1511 Siete Lagoas Minas Gerais Brazil

LA1512 Lago de Llopango El Salvador

LA1519 Vitarte Lima Peru

LA1540 Cali to Popayan Cauca Colombia

LA1542 Turrialba Costa Rica

LA1543 Upper Parana Brazil

LA1545 Becan Ruins Campeche Mexico

LA1546 Papantla Vera Cruz Mexico

LA1548 Fundo Liliana Junin Peru

LA1549 Chontabamba Pasco Peru

LA1569 Jalapa Vera Cruz Mexico

LA1574 Nana Lima Peru

LA1619 Pichanaki Junin Peru

LA1620 Castro Alves Bahia Brazil

LA1621 Rio Venados Hidalgo Mexico

LA1622 Lusaka Zambia

LA1623 Muna Yucatan Mexico

LA1632 Puerto Maldonado Madre de Dios Peru

LA1654 Tarapoto San Martin Peru

LA1655 Tarapoto San Martin Peru

LA1662 El Ejido Merida Venezuela

LA1667 Cali Cauca Colombia

LA1668 Acapulco Guerrero Mexico

LA1673 Nana Lima Peru

LA1701 Trujillo La Libertad Peru

LA1705 Sinaloa Mexico

LA1709 Desvio Yojoa Honduras

LA1710 Cariare Limon Costa Rica

LA1711 Zamorano Honduras

81

S. lycopersicum (L. esculentum var. cerasiforme)

LA1712 Pejibaye Costa Rica

LA1713 CATIE, Turrialba Costa Rica

LA1909 Quillabamba Cusco Peru

LA1953 La Curva Arequipa Peru

LA2076 Naranjitos Bolivia

LA2077 Paco, Coroica La Paz Bolivia

LA2078 Mosardas Rio Grande de Sol Brazil

LA2079 Maui: Kihei Hawaii USA

LA2080 Maui: Kihei Hawaii USA

LA2081 Maui: Kihei Hawaii USA

LA2082 Arenal Valley Honduras

LA2085 Kempton Park S. Africa

LA2095 La Cidra Loja Ecuador

LA2121 Yacuambi-Guadalupe Zamora-Chinchipe Ecuador

LA2122A Yacuambi-Guadalupe Zamora-Chinchipe Ecuador

LA2122B Yacuambi-Guadalupe Zamora-Chinchipe Ecuador

LA2122C Yacuambi-Guadalupe Zamora-Chinchipe Ecuador

LA2122D Yacuambi-Guadalupe Zamora-Chinchipe Ecuador

LA2123A La Saquea Zamora-Chinchipe Ecuador

LA2123B La Saquea Zamora-Chinchipe Ecuador

LA2126A El Dorado Zamora-Chinchipe Ecuador

LA2126B El Dorado Zamora-Chinchipe Ecuador

LA2126C El Dorado Zamora-Chinchipe Ecuador

LA2126D El Dorado Zamora-Chinchipe Ecuador

LA2127 Zumbi Zamora-Chinchipe Ecuador

LA2129 San Roque Zamora-Chinchipe Ecuador

LA2130 Gualaquiza Zamora-Chinchipe Ecuador

LA2131 Bomboiza Zamora-Chinchipe Ecuador

LA2135 Limon Santiago-Morona Ecuador

LA2136 Bella Union Santiago-Morona Ecuador

LA2137 Tayusa Santiago-Morona Ecuador

LA2138A Chinimpini Santiago-Morona Ecuador

LA2138B Chinimpini Santiago-Morona Ecuador

LA2139A Logrono Santiago-Morona Ecuador

LA2139B Logrono Santiago-Morona Ecuador

LA2140A Huambi Santiago-Morona Ecuador

LA2140B Huambi Santiago-Morona Ecuador

LA2140C Huambi Santiago-Morona Ecuador

LA2141 Rio Blanco Santiago-Morona Ecuador

LA2142 Cambanaca Santiago-Morona Ecuador

LA2143 Nuevo Rosario Santiago-Morona Ecuador

LA2177A San Ignacio Cajamarca Peru

82

S. lycopersicum (L. esculentum var. cerasiforme)

LA2177B San Ignacio Cajamarca Peru

LA2177C San Ignacio Cajamarca Peru

LA2177E San Ignacio Cajamarca Peru

LA2177F San Ignacio Cajamarca Peru

LA2205A Santa Rosa de Mirador San Martin Peru

LA2205B Santa Rosa de Mirador San Martin Peru

LA2308 San Francisco San Martin Peru

LA2312 Jumbilla #1 Amazonas Peru

LA2313 Jumbilla #2 Amazonas Peru

LA2392 Jakarta Indonesia

LA2393 Mercedes Canton Hoja Ancha Guanacaste Costa Rica

LA2394 San Rafael de Hoja Ancha Guanacaste Costa Rica

LA2402 Florianopolis Santa Catarina Brazil

LA2411 Yanamayo Puno Peru

LA2587 (4x, origin unknown)

LA2616 Naranjillo Huanuco Peru

LA2617 El Oropel Huanuco Peru

LA2618 Santa Lucia, Tulumayo Huanuco Peru

LA2619 Caseria San Augustin Loreto Peru

LA2620 La Divisoria Loreto Peru

LA2621 3 de Octubre Loreto Peru

LA2624 Umashbamba Cusco Peru

LA2625 Chilcachaca Cusco Peru

LA2626 Santa Ana Cusco Peru

LA2627 Pacchac, Chico Cusco Peru

LA2629 Echarate Cusco Peru

LA2630 Calzada Cusco Peru

LA2631 Chontachayoc Cusco Peru

LA2632 Maranura Cusco Peru

LA2633 Huayopata Cusco Peru

LA2635 Huayopata Cusco Peru

LA2636 Sicre Cusco Peru

LA2637 Sicre Cusco Peru

LA2640 Molinopata Apurimac Peru

LA2642 Molinopata Apurimac Peru

LA2643 Bella Vista Apurimac Peru

LA2660 San Ignacio de Moxos Beni Bolivia

LA2664 Yanahuana Puno Peru

LA2665 San Juan del Oro Puno Peru

LA2666 San Juan del Oro Puno Peru

LA2667 Pajchani Puno Peru

LA2668 Cruz Playa Puno Peru

LA2669 Huayvaruni #1 Puno Peru

83

S. lycopersicum (L. esculentum var. cerasiforme)

LA2670 Huayvaruni #2 Puno Peru

LA2671 San Juan del Oro, Escuela Puno Peru

LA2673 Chuntopata Puno Peru

LA2674 Huairurune Puno Peru

LA2675 Casahuiri Puno Peru

LA2683 Consuelo Cusco Peru

LA2684 Patria Cusco Peru

LA2685 Gavitana Madre de Dios Peru

LA2686 Yunguyo Madre de Dios Peru

LA2687 Mansilla Madre de Dios Peru

LA2688 Santa Cruz near Shintuyo #1 Madre de Dios Peru

LA2689 Santa Cruz near Shintuyo #2 Madre de Dios Peru

LA2690 Atalaya Cusco Peru

LA2691 Rio Pilcopata Cusco Peru

LA2692 Pilcopata #1 Cusco Peru

LA2693 Pilcopata #2 Cusco Peru

LA2694 Aguasantas Cusco Peru

LA2696 El Paramillo, La Union Valle Colombia

LA2697 Mata de Cana, El Dovio Valle Colombia

LA2698 La Esperanza de Belgica Valle Colombia

LA2700 Aoti, Satipo Junin Peru

LA2702 Kandy #1 Sri Lanka

LA2709 Bidadi, Bangalore Karnataka India

LA2710 Porto Firme Brazil

LA2782 El Volcan #1 - Pajarito Antioquia Colombia

LA2783 El Volcan #2 - Titiribi Antioquia Colombia

LA2784 La Queronte Antioquia Colombia

LA2785 El Bosque Antioquia Colombia

LA2786 Andes #1 Antioquia Colombia

LA2787 Andes #2 Antioquia Colombia

LA2789 Canaveral Antioquia Colombia

LA2790 Buenos Aires Antioquia Colombia

LA2791 Rio Frio Antioquia Colombia

LA2792 Tamesis Antioquia Colombia

LA2793 La Mesa Antioquia Colombia

LA2794 El Libano Antioquia Colombia

LA2795 Camilo Antioquia Colombia

LA2807 Taypiplaya Yungas Bolivia

LA2811 Cerro Huayrapampa Apurimac Peru

LA2814 Ccascani, Sandia Puno Peru

LA2841 Chinuna Amazonas Peru

LA2842 Santa Rita San Martin Peru

LA2843 Moyobamba mercado San Martin Peru

84

S. lycopersicum (L. esculentum var. cerasiforme)

LA2844 Shanhao San Martin Peru

LA2845 Mercado Moyobamba San Martin Peru

LA2871 Chamaca Sud Yungas Bolivia

LA2873 Lote Pablo Luna #2 Sud Yungas Bolivia

LA2874 Playa Ancha Sud Yungas Bolivia

LA2933 Jipijapa Manabi Ecuador

LA2977 Belen Beni Bolivia

LA2978 Belen Beni Bolivia

LA3135 Pinal del Jigue Holguin Cuba

LA3136 Arroyo Rico Holguin Cuba

LA3137 Pinares de Mayari Holguin Cuba

LA3138 El Quemada Holguin Cuba

LA3139 San Pedro de Cananova Holguin Cuba

LA3140 Los Platanos Holguin Cuba

LA3141 Guira de Melena La Habana Cuba

LA3162 N of Copan Honduras

LA3452 CATIE, Turrialba Turrialba Costa Rica

LA3623 Tablones Manabi Ecuador

LA3633 Botanical garden Ghana

LA3652 Matara Apurimac Peru

LA3842 El Limon, Maracay Araguay Venezuela

LA3843 El Limon, Maracay Aragua Venezuela

LA3844 Algarrobito Guarico Venezuela

LA4133 Makapuu Beach, Oahu Hawaii USA

LA4352 Bamoa Sinaloa Mexico

LA4353 Guasave Sinaloa Mexico

S. neorickii (L. parviflorum)

LA0247 Chavinillo Huanuco Peru

LA0735 Huariaca Huanuco Peru

LA1319 Abancay Apurimac Peru

LA1321 Curahuasi Apurimac Peru

LA1322 Limatambo Cusco Peru

LA1326 Rio Pachachaca Apurimac Peru

LA1329 Yaca Apurimac Peru

LA1626A Mouth of Rio Rupac Ancash Peru

LA1716 Huancabamba Piura Peru

LA2072 Huanuco Huanuco Peru

LA2073 Huanuco, N of San Rafael Huanuco Peru

LA2074 Huanuco Huanuco Peru

LA2075 Huanuco Huanuco Peru

LA2113 La Toma Loja Ecuador

LA2133 Ona Azuay Ecuador

85

S. neorickii (L. parviflorum)

LA2190 Tialango Amazonas Peru

LA2191 Campamento Ingenio Amazonas Peru

LA2192 Pedro Ruiz Amazonas Peru

LA2193 Churuja Amazonas Peru

LA2194 Chachapoyas West Amazonas Peru

LA2195 Caclic Amazonas Peru

LA2197 Luya Amazonas Peru

LA2198 Chachapoyas East Amazonas Peru

LA2200 Choipiaco Amazonas Peru

LA2201 Pipus Amazonas Peru

LA2202 Tingobamba Amazonas Peru

LA2315 Sargento Amazonas Peru

LA2317 Zuta Amazonas Peru

LA2318 Lima Tambo Amazonas Peru

LA2319 Chirico Amazonas Peru

LA2325 Above Balsas Amazonas Peru

LA2403 Wandobamba Huanuco Peru

LA2613 Matichico-San Rafael Huanuco Peru

LA2614 San Rafael Huanuco Peru

LA2615 Ayancocho Huanuco Peru

LA2639A Puente Cunyac Apurimac Peru

LA2641 Nacchera Apurimac Peru

LA2727 Ona Azuay Ecuador

LA2847 Suyubamba Amazonas Peru

LA2848 Pedro Ruiz Amazonas Peru

LA2862 Saraguro-Cuenca Azuay Ecuador

LA2865 Rio Leon Azuay Ecuador

LA2913 Uchucyaco - Hujainillo Huanuco Peru

LA3651 Matara Apurimac Peru

LA3655 Casinchigua to Chacoche Apurimac Peru

LA3657 Casinchigua to Pichirhua Apurimac Peru

LA3660 Murashaya Apurimac Peru

LA3793 Huariaca to San Rafael Huanuco Peru

LA4020 Gonozabal Loja Ecuador

LA4021 Guancarcucho Azuay Ecuador

LA4022 Pueblo Nuevo Azuay Ecuador

LA4023 Paute Azuay Ecuador

S. ochranthum

LA2118 San Lucas Loja Ecuador

LA2160 Acunac Cajamarca Peru

LA2161 Cruz Roja Cajamarca Peru

LA2162 Yatun Cajamarca Peru

86

S. ochranthum

LA2166 Pacopampa Cajamarca Peru

LA2203 Pomacochas San Martin Peru

LA2682 Chinchaypujio Cusco Peru

LA3649 Curpahuasi-Pacaipampa Apurimac Peru

LA3650 Choquemaray Apurimac Peru

S. pennellii (L. pennellii, L. pennellii var. puberulum)

LA0716 Atico Arequipa Peru

LA0750 Ica to Nazca Ica Peru

LA0751 Sisacaya Lima Peru

LA1272 Pisaquera Lima Peru

LA1273 Cayan Lima Peru

LA1275 Quilca road junction Lima Peru

LA1277 Trapiche Lima Peru

LA1282 Sisacaya Lima Peru

LA1297 Pucara Lima Peru

LA1299 Santa Rosa de Quives Lima Peru

LA1302 Quita Sol Ica Peru

LA1303 Pampano Huancavelica Peru

LA1340 Capillucas Lima Peru

LA1356 Moro Ancash Peru

LA1367 Santa Eulalia Lima Peru

LA1376 Sayan Lima Peru

LA1515 Sayan to Churin Lima Peru

LA1522 Quintay Lima Peru

LA1649 Molina Ica Peru

LA1656 Marca to Chincha Ica Peru

LA1657 Buena Vista to Yautan Ancash Peru

LA1674 Toparilla Canyon Lima Peru

LA1693 Quebrada Machurango Lima Peru

LA1724 La Quinga Ica Peru

LA1732 Rio San Juan Huancavelica Peru

LA1733 Rio Canete Lima Peru

LA1734 Rio Canete Lima Peru

LA1735 Rio Canete Lima Peru

LA1809 El Horador (playa) Piura Peru

LA1911 Locari Ica Peru

LA1912 Cerro Locari Ica Peru

LA1920 Cachiruma (Rio Grande) Ayacucho Peru

LA1926 Agua Perdida (Rio Ingenio) Ica Peru

LA1940 Rio Atico, Km 26 Arequipa Peru

LA1941 Rio Atico, Km 41 Arequipa Peru

LA1942 Rio Atico, Km 54 Arequipa Peru

87

S. pennellii (L. pennellii, L. pennellii var. puberulum)

LA1943 Rio Atico, Km 61 Arequipa Peru

LA1946 Caraveli Arequipa Peru

LA2560 Santa to Huaraz Ancash Peru

LA2580 Valle de Casma Ancash Peru

LA2657 Bayovar Piura Peru

LA2963 Acoy Arequipa Peru

LA3635 Omas Lima Peru

LA3665 Ica to Nazca (Rio Santa Cruz) Ica Peru

LA3778 Palpa to Nazca Ica Peru

LA3788 Rio Atico, Km 10 Arequipa Peru

LA3789 Rio Atico, Km 26 Arequipa Peru

LA3791 Caraveli Arequipa Peru

S. peruvianum (L. peruvianum)

LA0098 Chilca Lima Peru

LA0111 Supe Lima Peru

LA0153 Culebras Ancash Peru

LA0370 Hacienda Huampani Lima Peru

LA0371 Supe Lima Peru

LA0372 Culebras #1 Ancash Peru

LA0374 Culebras #2 Ancash Peru

LA0445 Chincha #2 Ica Peru

LA0446 Atiquipa Arequipa Peru

LA0448 Chala Arequipa Peru

LA0453 Yura Arequipa Peru

LA0454 Tambo Arequipa Peru

LA0455 Tambo Arequipa Peru

LA0462 Sobraya Tarapaca Chile

LA0464 Hacienda Rosario Tarapaca Chile

LA0752 Sisacaya Lima Peru

LA1161 Huachipa Lima Peru

LA1270 Pisiquillo Lima Peru

LA1278 Trapiche Lima Peru

LA1300 Santa Rosa de Quives Lima Peru

LA1333 Loma Camana Arequipa Peru

LA1336 Atico Arequipa Peru

LA1337 Atiquipa Arequipa Peru

LA1368 San Jose de Palla Lima Peru

LA1369 San Geronimo Lima Peru

LA1474 Lomas de Camana Arequipa Peru

LA1475 Fundo 'Los Anitos', Barranca Lima Peru

LA1513 Atiquipa Arequipa Peru

LA1517 Irrigacion Santa Rosa Lima Peru

88

S. peruvianum (L. peruvianum)

LA1537 Azapa Valley Tarapaca Chile

LA1556 Hacienda Higuereta Lima Peru

LA1616 La Rinconada Lima Peru

LA1675 Toparilla Canyon Lima Peru

LA1692 Putinza Lima Peru

LA1913 Tinguiayog Ica Peru

LA1929 La Yapana (Rio Ingenio) Ica Peru

LA1935 Lomas de Atiquipa Arequipa Peru

LA1947 Puerto Atico Arequipa Peru

LA1949 Las Calaveritas Arequipa Peru

LA1951 Ocona Arequipa Peru

LA1954 Mollendo Arequipa Peru

LA1955 Matarani Arequipa Peru

LA1975 Desvio Santo Domingo Lima Peru

LA1977 Orcocoto Lima Peru

LA1989 (self-fertile selection, origin unknown)

LA2573 Valle de Casma Ancash Peru

LA2581 Chacarilla (4x) Tarapaca Chile

LA2732 Moquella Tarapaca Chile

LA2742 Camarones-Guancarane Tarapaca Chile

LA2744 Sobraya Tarapaca Chile

LA2745 Pan de Azucar Tarapaca Chile

LA2770 Lluta Tarapaca Chile

LA2834 Hacienda Asiento Ica Peru

LA2955B Quistagama Tarapaca Chile

LA2959 Chaca to Caleta Vitor Tarapaca Chile

LA2964 Quebrada de Burros Tacna Peru

LA3218 Quebrada Guerrero (Islay) Arequipa Peru

LA3220 Cocachacra Arequipa Peru

LA3636 Coayllo Lima Peru

LA3640 Mexico City Mexico

LA3781 Quebrada Oscollo (Atico) Arequipa Peru

LA3783 Rio Chaparra Arequipa Peru

LA3787 Alta Chaparra Arequipa Peru

LA3790 Caraveli Arequipa Peru

LA3795 Alta Fortaleza Ancash Peru

LA3797 Anca, Marca (Rio Fortaleza) Ancash Peru

LA3799 Río Pativilca Ancash Peru

LA3853 Mollepampa La Libertad Peru

LA3858 Canta Lima Peru

LA3900 (CMV tolerant selection)

LA4125 Camina Tarapaca Chile

LA4128 Pachica (Rio Camarones) Tarapaca Chile

89

S. peruvianum (L. peruvianum)

LA4317 Rio Lluta, desembocadura Tarapaca Chile

LA4318 Sora - Molinos, Rio Lluta Tarapaca Chile

LA4325 Caleta Vitor Tarapaca Chile

LA4328 Pachica, Rio Camarones Tarapaca Chile

LA4445 Azapa Valley, 27 km from Arica Tarapaca Chile

LA4446 Azapa Valley, Km 37 from Arica Tarapaca Chile

LA4447 Azapa Valley, Km 27 and Km 37 from Arica Tarapaca Chile

S. pimpinellifolium (L. pimpinellifolium)

LA0100 La Cantuta (Rimac Valley) Lima Peru

LA0114 Pacasmayo La Libertad Peru

LA0121 Trujillo La Libertad Peru

LA0122 Poroto La Libertad Peru

LA0369 La Cantuta (Rimac Valley) Lima Peru

LA0373 Culebras #1 Ancash Peru

LA0375 Culebras #2 Ancash Peru

LA0376 Hacienda Chiclin La Libertad Peru

LA0381 Pongo La Libertad Peru

LA0391 Magdalena (Rio Jequetepeque) Cajamarca Peru

LA0397 Hacienda Tuman Lambayeque Peru

LA0398 Hacienda Carrizal Cajamarca Peru

LA0400 Hacienda Buenos Aires Piura Peru

LA0411 Pichilingue Los Rios Ecuador

LA0412 Pichilingue Los Rios Ecuador

LA0413 Cerecita Guayas Ecuador

LA0417 Puna Guayas Ecuador

LA0418 Daule Guayas Ecuador

LA0420 El Empalme Guayas Ecuador

LA0442 Sechin Ancash Peru

LA0443 Pichilingue Los Rios Ecuador

LA0480 Hacienda Santa Inez Ica Peru

LA0722 Trujillo La Libertad Peru

LA0753 Lurin Lima Peru

LA1236 Tinelandia, Santo Domingo Pichincha Ecuador

LA1237 Atacames Esmeraldas Ecuador

LA1242 Los Sapos Guayas Ecuador

LA1243 Co-op Carmela Guayas Ecuador

LA1245 Santa Rosa El Oro Ecuador

LA1246 La Toma Loja Ecuador

LA1248 Hacienda Monterrey Loja Ecuador

LA1256 Naranjal Guayas Ecuador

LA1257 Las Mercedes Guayas Ecuador

LA1258 Voluntario de Dios Guayas Ecuador

90

S. pimpinellifolium (L. pimpinellifolium)

LA1259 Catarama Los Rios Ecuador

LA1260 Pueblo Viejo Los Rios Ecuador

LA1261 Babahoyo Los Rios Ecuador

LA1262 Milagro Empalme Guayas Ecuador

LA1263 Barranco Chico Guayas Ecuador

LA1269 Pisiquillo Lima Peru

LA1279 Cieneguilla Lima Peru

LA1280 Chontay Lima Peru

LA1301 Hacienda San Ignacio Ica Peru

LA1332 Nazca Ica Peru

LA1335 Pescaderos Arequipa Peru

LA1341 Huampani Lima Peru

LA1342 Casma Ancash Peru

LA1343 Puente Chao La Libertad Peru

LA1344 Laredo La Libertad Peru

LA1345 Samne La Libertad Peru

LA1348 Pacasmayo La Libertad Peru

LA1349 Cuculi Lambayeque Peru

LA1355 Nepena Ancash Peru

LA1357 Jimbe Ancash Peru

LA1359 La Crau Ancash Peru

LA1370 San Jose de Palla Lima Peru

LA1371 Santa Eulalia Lima Peru

LA1374 Ingenio Ica Peru

LA1375 San Vicente de Canete Lima Peru

LA1380 Chanchape Piura Peru

LA1381 Naupe Lambayeque Peru

LA1382 Chachapoyas to Balsas Amazonas Peru

LA1383 Chachapoyas to Bagua Amazonas Peru

LA1384 Quebrada Parca Lima Peru

LA1416 Las Delicias Pichincha Ecuador

LA1428 La Estancilla Manabi Ecuador

LA1466 Chongoyape Lambayeque Peru

LA1469 El Pilar, Olmos Lambayeque Peru

LA1470 Motupe to Desvio Olmos-Bagua Lambayeque Peru

LA1471 Motupe to Jayanca Lambayeque Peru

LA1472 Quebrada Topara Lima Peru

LA1478 Santo Tome (Pabur) Piura Peru

LA1514 Sayan to Churin Lima Peru

LA1520 Sayan to Churin Lima Peru

LA1521 El Pinon, Asia Lima Peru

LA1547 Chota to El Angel Carchi Ecuador

LA1561 San Eusebio Lima Peru

91

S. pimpinellifolium (L. pimpinellifolium)

LA1562 Cieneguilla Lima Peru

LA1571 San Jose de Palle Lima Peru

LA1572 Hacienda Huampani Lima Peru

LA1573 Nana Lima Peru

LA1575 Huaycan Lima Peru

LA1576 Manchay Alta Lima Peru

LA1577 Cartavio La Libertad Peru

LA1578 Santa Marta La Libertad Peru

LA1579 Colegio Punto Cuatro #1 Lambayeque Peru

LA1580 Colegio Punto Cuatro #2 Lambayeque Peru

LA1581 Punto Cuatro Lambayeque Peru

LA1582 Motupe Lambayeque Peru

LA1583 Tierra de la Vieja Lambayeque Peru

LA1584 Jayanca to La Vina Lambayeque Peru

LA1585 Cuculi Lambayeque Peru

LA1586 Zana, San Nicolas La Libertad Peru

LA1587 San Pedro de Lloc La Libertad Peru

LA1588 Laredo to Barraza La Libertad Peru

LA1589 Viru to Galunga La Libertad Peru

LA1590 Viru to Tomaval La Libertad Peru

LA1591 Ascope La Libertad Peru

LA1592 Moche La Libertad Peru

LA1593 Puente Chao La Libertad Peru

LA1594 Cerro Sechin Ancash Peru

LA1595 Nepena to Samanco Ancash Peru

LA1596 Santa to La Rinconada Ancash Peru

LA1597 Rio Casma Ancash Peru

LA1598 Culebras to La Victoria Ancash Peru

LA1599 Huarmey Ancash Peru

LA1600 Las Zorras, Huarmey Ancash Peru

LA1601 La Providencia Lima Peru

LA1602 Rio Chillon to Punchauca Lima Peru

LA1603 Quilca Lima Peru

LA1604 Horcon Lima Peru

LA1605 Canete - San Antonio Lima Peru

LA1606 Tambo de Mora Ica Peru

LA1607 Canete - La Victoria Lima Peru

LA1608 Canete - San Luis Lima Peru

LA1610 Asia - El Pinon Lima Peru

LA1611 Rio Mala Lima Peru

LA1612 Rio Chilca Lima Peru

LA1613 Santa Eusebia Lima Peru

LA1614 Pampa Chumbes Lima Peru

92

S. pimpinellifolium (L. pimpinellifolium)

LA1615 Piura to Simbala Piura Peru

LA1617 Tumbes South Tumbes Peru

LA1618 Tumbes North Tumbes Peru

LA1628 Huanchaco La Libertad Peru

LA1629 Barrancos de Miraflores Lima Peru

LA1630 Fundo La Palma Ica Peru

LA1631 Planta Envasadora San Fernando (Moche) La Libertad Peru

LA1633 Co-op Huayna Capac Ica Peru

LA1634 Fundo Bogotalla #1 Ica Peru

LA1635 Fundo Bogotalla #2 Ica Peru

LA1636 Laran Ica Peru

LA1637 La Calera Ica Peru

LA1638 Fundo El Portillo Lima Peru

LA1645 Banos de Miraflores Lima Peru

LA1651 Vivero, La Molina Lima Peru

LA1652 Cieneguilla Lima Peru

LA1659 Pariacoto Ancash Peru

LA1660 Yautan to Pariacoto Ancash Peru

LA1661 Esquina de Asia Lima Peru

LA1670 Rio Sama Tacna Peru

LA1676 Fundo Huadquina, Topara Ica Peru

LA1678 San Juan Lucumo de Topara Ica Peru

LA1679 Tambo de Mora Ica Peru

LA1680 La Encanada Lima Peru

LA1682 Montalban - San Vicente Lima Peru

LA1683 Miramar Piura Peru

LA1684 Chulucanas Piura Peru

LA1685 Marcavelica Piura Peru

LA1686 Valle Hermosa #1 Piura Peru

LA1687 Valle Hermoso #2 Piura Peru

LA1688 Pedregal Piura Peru

LA1689 Castilla #1 Piura Peru

LA1690 Castilla #2 Piura Peru

LA1697 Hacienda Quiroz, Santa Anita Lima Peru

LA1719 E of Arenillas El Oro Ecuador

LA1720 Yautan Ancash Peru

LA1728 Rio San Juan Ica Peru

LA1729 Rio San Juan Ica Peru

LA1742 Olmos-Marquina Lambayeque Peru

LA1781 Bahia de Caraquez Manabi Ecuador

LA1921 Majarena Ica Peru

LA1923 Cabildo Ica Peru

LA1924 Piedras Gordas Ica Peru

93

S. pimpinellifolium (L. pimpinellifolium)

LA1925 Pangaravi Ica Peru

LA1933 Jaqui Arequipa Peru

LA1936 Huancalpa Arequipa Peru

LA1950 Pescadores Arequipa Peru

LA1987 Viru-Fundo Luis Enrique La Libertad Peru

LA1992 Pichicato Lima Peru

LA1993 Chicama Valley? Lima Peru

LA2093 La Union El Oro Ecuador

LA2096 Playa Loja Ecuador

LA2097 Macara Loja Ecuador

LA2102 El Lucero Loja Ecuador

LA2112 Hacienda Monterrey Loja Ecuador

LA2145 Juan Montalvo Los Rios Ecuador

LA2146 Hacienda Limoncarro La Libertad Peru

LA2147 Yube Cajamarca Peru

LA2149 Puente Muyuno Cajamarca Peru

LA2170 Pai Pai Cajamarca Peru

LA2173 Cruz de Huayquillo Cajamarca Peru

LA2176 Timbaruca Cajamarca Peru

LA2178 Tororume Cajamarca Peru

LA2179 Tamboripa - La Manga Cajamarca Peru

LA2180 La Coipa Cajamarca Peru

LA2181 Balsa Huaico Cajamarca Peru

LA2182 Cumba Amazonas Peru

LA2183 Corral Quemado Amazonas Peru

LA2184 Bagua Amazonas Peru

LA2186 El Salao Amazonas Peru

LA2187 La Caldera Amazonas Peru

LA2188 Machugal #1 Amazonas Peru

LA2189 Machugal #2 Amazonas Peru

LA2335 (4x)

LA2340 (4x)

LA2345 (doubled haploid, origin unknown)

LA2346 (doubled haploid, origin unknown)

LA2347 (doubled haploid, origin unknown)

LA2348 Trujillo La Libertad Peru

LA2389 Tembladera Cajamarca Peru

LA2390 Chungal Cajamarca Peru

LA2391 Chungal to Monte Grande Cajamarca Peru

LA2401 Moxeque Ancash Peru

LA2412 Fundo Don Javier, Chilca Lima Peru

LA2533 Lomas de Latillo Lima Peru

LA2576 Valle de Casma Ancash Peru

94

S. pimpinellifolium (L. pimpinellifolium)

LA2578 Tuturo Ancash Peru

LA2585 (4x, origin unknown)

LA2628 Echarate Cusco Peru

LA2645 Desvio Chulucanas-Morropon Piura Peru

LA2646 Chalaco Piura Peru

LA2647 Morropon-Chalaco Piura Peru

LA2652 Sullana Piura Peru

LA2653 San Francisco de Chocon Querecotillo Piura Peru

LA2655 La Huaca to Sullana Piura Peru

LA2656 Suarez Tumbes Peru

LA2659 Castilla, Univ. Nac. de Piura Piura Peru

LA2718 Chilca Lima Peru

LA2725 Tambo Colorado Ica Peru

LA2805 cv. Indehiscent Currant

LA2831 Rio Nazca Ica Peru

LA2832 Chicchi Tara Ica Peru

LA2833 Hacienda Asiento Ica Peru

LA2836 Fundo Pongo Ica Peru

LA2839 Tialango Amazonas Peru

LA2840 San Hilarion de Tomaque Amazonas Peru

LA2850 Santa Rosa, Manta Manabi Ecuador

LA2851 La Carcel de Montecristo Manabi Ecuador

LA2852 Cirsto Rey de Charapoto Manabi Ecuador

LA2853 Experiment Station, Portoviejo-INIAP Manabi Ecuador

LA2854 Jipijapa Manabi Ecuador

LA2857 Isabela: Puerto Villamil Galapagos Islands Ecuador

LA2866 Via a Amaluza Loja Ecuador

LA2914A Urb. La Castellana, Surco Lima Peru

LA2914B La Castellana, Surco Lima Peru

LA2915 El Remanso de Olmos Lambayeque Peru

LA2934 Carabayllo Lima Peru

LA2966 La Molina Lima Peru

LA2974 Huaca del Sol La Libertad Peru

LA2982 Chilca #1 Lima Peru

LA2983 Chilca #2 Lima Peru

LA3123 Santa Cruz: summit Galapagos Islands Ecuador

LA3158 Los Mochis Sinaloa Mexico

LA3159 Los Mochis Sinaloa Mexico

LA3160 Los Mochis Sinaloa Mexico

LA3161 Los Mochis Sinaloa Mexico

LA3468 La Molina Vieja Lima Peru

LA3634 Santa Rosa de Asia Lima Peru

LA3638 Ccatac Lima Peru

95

S. pimpinellifolium (L. pimpinellifolium)

LA3798 Río Pativilca Ancash Peru

LA3803 Pacanguilla La Libertad Peru

LA3852 Atinchik, Pachacamac Lima Peru

LA3859 TYLCV resistant selection „hirsute‟

LA3910 Near tortoise preserve, Santa Cruz Galapagos Islands Ecuador

LA4027 Olmos-Jaen Road Lambayeque Peru

LA4138 El Corregidor, La Molina Lima Peru

S. sitiens (S. rickii)

LA1974 Chuquicamata Antofagasta Chile

LA2876 Chuquicamata Antofagasta Chile

LA2877 El Crucero Antofagasta Chile

LA2878 Mina La Exotica Antofagasta Chile

LA2885 Caracoles Antofagasta Chile

LA4105 Mina La Escondida Antofagasta Chile

LA4110 Mina San Juan Antofagasta Chile

LA4112 Aguada Limon Verde Antofagasta Chile

LA4113 Estacion Cere Antofagasta Chile

LA4114 Pampa Carbonatera Antofagasta Chile

LA4115 Quebrada desde Cerro Oeste de Paqui Antofagasta Chile

LA4116 Quebrada de Paqui Antofagasta Chile

LA4331 Cerro Quimal Antofagasta Chile

96

Appendix. Wild species core collections. The accession numbers included in the core subsets for each species are listed below. In addition, this table lists accessions in the SolCAP core which are derived from the TGRC (wild species only). The „species sampler‟ subset includes 2-3 accessions from each species group.

S. arcanum

LA0441

LA1346

LA1360

LA1626

LA1708

LA1984

LA2152

LA2163

LA2172

LA2185

LA2326

LA2328

LA2553

S. cheesmaniae

LA0428

LA0429

LA0531

LA1039

LA1041

LA1406

LA1407

LA1409

LA1412

LA1450

S. chilense

LA1930

LA1932

LA1958

LA1960

LA1963

LA1967

LA1969

LA1971

LA2748

LA2750

LA2753

LA2759

LA2765

S. chilense

LA2771

LA2778

LA2880

LA2884

LA2930

LA2946

LA3114

S. chmielewskii

LA1028

LA1306

LA1316

LA1317

LA1325

LA1330

LA2663

LA2677

LA2680

LA2695

LA0103

S. corneliomulleri

LA0107

LA0444

LA1292

LA1305

LA1331

LA1339

LA1647

LA1677

LA1910

LA1937

LA1945

LA1973

S. galapagense

LA0317

LA0438

LA0483

LA0526

LA1136

97

S. galapagense

LA1137

LA1141

LA1401

LA1410

S. habrochaites

LA0407

LA1223

LA1266

LA1347

LA1353

LA1361

LA1363

LA1559

LA1624

LA1718

LA1721

LA1731

LA1753

LA1777

LA1918

LA1928

LA2098

LA2103

LA2109

LA2119

LA2128

LA2155

LA2158

LA2167

LA2174

LA2204

LA2329

LA2409

LA2650

LA2864

S. huaylasense

LA1364

LA1365

LA1982

LA2808

S. lyc. cerasiforme

LA0292

LA1204

S. lyc. cerasiforme

LA1206

LA1228

LA1231

LA1268

LA1286

LA1307

LA1314

LA1320

LA1323

LA1338

LA1385

LA1388

LA1420

LA1425

LA1429

LA1453

LA1456

LA1461

LA1464

LA1482

LA1483

LA1509

LA1511

LA1542

LA1543

LA1620

LA1622

LA2078

LA2095

LA2131

LA2138A

LA2308

LA2392

LA2402

LA2621

LA2670

LA2675

LA2688

LA2709

LA2710

LA2783

LA2845

LA2871

98

S. lyc. cerasiforme

LA4133

LA0247

S. neorickii

LA1319

LA1322

LA1626A

LA1716

LA2113

LA2133

LA2190

LA2198

LA2319

LA2325

S. pennellii

LA0716

LA0751

LA1272

LA1277

LA1356

LA1367

LA1376

LA1656

LA1674

LA1724

LA1732

LA1733

LA1926

LA1946

LA2580

LA2963

S. peruvianum

LA0153

LA0446

LA0752

LA1274

LA1336

LA1474

LA1954

LA2732

LA2744

S. pimpinellifolium

LA0373

LA0400

S. pimpinellifolium

LA0411

LA0417

LA0442

LA1237

LA1245

LA1246

LA1261

LA1279

LA1301

LA1335

LA1371

LA1375

LA1478

LA1521

LA1547

LA1576

LA1578

LA1582

LA1584

LA1586

LA1590

LA1593

LA1599

LA1602

LA1606

LA1617

LA1659

LA1683

LA1689

LA1924

LA1936

LA2102

LA2173

LA2181

LA2183

LA2401

LA2533

LA2852

SolCAP

LA0166

LA0317

LA0373

LA0407

99

SolCAP

LA0422

LA0438

LA0446

LA0716

LA0722

LA1028

LA1037

LA1141

LA1208

LA1237

LA1246

LA1269

LA1272

LA1274

LA1282

LA1283

LA1290

LA1301

LA1314

LA1322

LA1331

LA1338

LA1340

LA1346

LA1406

LA1455

LA1457

LA1464

LA1478

LA1512

LA1542

LA1545

LA1547

LA1549

LA1569

LA1578

LA1582

LA1589

LA1617

LA1620

LA1621

LA1623

LA1632

SolCAP

LA1654

LA1656

LA1668

LA1674

LA1701

LA1712

LA1732

LA1777

LA1809

LA1912

LA1926

LA1930

LA1941

LA1946

LA1953

LA1963

LA1973

LA2076

LA2077

LA2078

LA2093

LA2099

LA2126A

LA2131

LA2135

LA2137

LA2163

LA2181

LA2184

LA2185

LA2190

LA2308

LA2312

LA2411

LA2533

LA2560

LA2561

LA2626

LA2632

LA2633

LA2660

LA2663

LA2664

100

SolCAP

LA2675

LA2744

LA2779

LA2788

LA2792

LA2852

LA2880

LA2930

LA2932

LA2951

LA3136

LA3137

LA3650

LA3795

LA4331

Species Sampler

LA0528

LA0716

LA0722

LA1037

LA1223

LA1226

LA1274

LA1293

LA1326

LA1589

LA1777

LA1926

LA1932

LA1982

LA2150

LA2663

LA2884

LA2930

LA3661

101

Membership List Aarden, Harriette Monsanto Holland BV, Dept. Tomato Breeding.

Leeuwenhoekweg 52, CZ Bergschenhoek, 2661 harriette.aarden@monsanto.com

Alger, Hillary Johnny's Selected Seeds, USA halger@johnnyseeds.com ARC Veg and Orn Plant Inst. Atanassiva, Bistra Institute of Genetics, Prof. D. Kostov, BAS, Plovdivsko

Choss 13km, Sofia, BULGARIA, 1113 bistra_a@yahoo.com Augustine, Jim BHN Research/ BHN Seed, PO Box 3267, Immokalee, Fl, USA,

34143 jaugustine@bhnseed.com Beck Bunn, Teresa Monsanto/Seminis, 37437 State Hwy 16, Woodland, CA,

USA, 95695 teresa.beck.bunn@seminis.com Beckles, Diane M. University of Cal- Davis, Plant Sciences- MS3, One Shields Ave,

Davis, CA, USA, 95616 dmbeckles@ucdavis.edu Buonfiglioli, Carlo Della Rimembranze nr. 6A, San Lazzaro di Savena, Bologna, ITALY, 40068 red@prorainbow.com Burdick, Allan 3000 Woodkirk Dr., Columbia, MO, USA, 65203 allanburdick@pocketmail.com California Tomato Research Institute, Inc. Library 18650 E. Lone Tree Rd., Escalon,

CA, USA, 95320-9759 Carli, Stefano Nunhems Italy, via Ghiarone 2, S.Agata, Bolongnese, ITALY,

40019 stefano.carli@nunhems.com Carrijo Iedo, Valentim Rua Joao Angelo do Pinho 77, Apto 102, Betim, MG, BRAZIL, 32.510-040 iedovc@uai.com.br Chen, Dei Wei Bucolic Seeds Co. Ltd., P.O. 2-39, Tantzu, Taichung Co., TAIWAN, 427 bucolic.seeds@msa.hinet.net Chetelat, Roger University of California, Dept of Veg Crops, One Shields Ave,

Davis, CA, USA, 95616-8746 trchetelat@ucdavis.edu . Cornell University, Albert R Mann Library, Serials Unit/Acquisition Div, Ithaca, NY, USA,

14853

102

Coulibaly, Sylvaine Nunhems USA, 7087 E. Peltier Rd, Acampo, CA, USA, 95220 sylvaine.coulibaly@nunhems.com Cuartero, Jesus E.E. LaMayora- CSIC, Plant Breeding Dept., Algarrobo-Costa, Malaga, SPAIN, 29760 rfern@eelm.csic.es deHoop, Simon Jan East West Seed Co. Ltd, PO Box 3, Bang Bua Thong, Nonthaburi, THAILAND, 11110 simon.dehoop@eastwestseed.com Dick, Jim Tomato Solutions, 23264 Mull Rd, Chatham, Ontario, CANADA, N7M 5J4 jimdick@netrover.com Fernandez-Munoz, Rafael E.E. LaMayora- CSIC, Plant Breeding Dept,

Algarrobo-Costa, Malaga, SPAIN, 29750 rfern@eelm.csic.es Fisher, Dr. Dave K. Fisher Farms, 48244 Wesley Chapel Rd, Richfield, NC,

USA, 28137 fisherfarms1933@earthlink.net Foolad, Majid Penn State University, Dept. of Horticulture, 102 Tyson Blvd.,

University Park, PA, USA, 16802 mrf5@psu.edu Fowler, C. Wayne, 2840 70th St SW, Naples, FL, USA, 34105 c_w_fowler@msn.com Frank A. Lee Library, NYS Agriculture Experimental Station, 630 W. North St, Geneva,

NY, USA, 14456-1371 Georgiev, Hristo Atanasov Urbanizacion, COSTA JARDIN,

Residencial LA GRACIOSA, C/. LA NUBE 20, Telde, SPAIN, 35215 eliedi1003@yahoo.es

Gorin, Anthony Technisem, 7 au du Gargliano, Zac des Gatines, FRANCE, 91600 anthony.gorin@technisem.com Grazzini, Rick GardenGenetics LLC, 131 Mendels Way, Bellefonte, PA, USA, 16823 rick@gardengenetics.com Hanson, Peter AVRDC, PO Box 42, Shanhua, Tainan, TAIWAN,

REPUBLIC of CHINA, 741 hansp@netra.avrdc.org.tw Hayashi, Masako Yaguchi Asahi Industries, Biol.Engineering Lab, 222 Watarase,

Kamikawa, Kodama-gun, Saitama-ken, JAPAN, 367-0394 m.hayashi@asahi-kg.co.jp

Hernandez, Rogelio Harris Moran, 2092 Mission Drive, Naples, FL, USA, 34109 r.hernandez@harrismoran.com

103

Hoogstraten, Jaap Seminis Veg Seeds, Postbus 97, 6700 AB Wageningen, THE NETHERLANDS jaap.hoogstraten@seminis.com

Hotzev, Amit AB-SEEDS, ltd., P.O. Box 1, Teradion Ind. Zone, D.N. MISGAV, ISRAEL, 20179 amit.hotzev@ab-seeds.com Hutton, Sam University of Florida, Gulf Coast Research and Education Center, 14625 County Rd 672, Wimuama, FL, USA, 33598 sfhutton@ufl.edu Ignatova, Svetlana Box 15, Moscow E-215, RUSSIA, 105215 sril@bk.ru Inai, Shuji Nippon Del Monte Corp., Research and Development, 3748 Shimizu-Cho Numata-shi, Gunma-ken, JAPAN, 378-0016 sinai@delmonte.co.jp Indian Institute of Horticultural Research, Bangalore, INDIA Jahrmann, Torben Semillas Fito, Centre de biotecnologia, Riera d/Agell, 11, Cabrera de Mar, Barcelona, SPAIN, 8349 tjahrmann@semillasfito.com Johnston, Rob Johnny's Selected Seeds, 955 Benton Ave, Winslow, ME, USA, 4901 rjohnston@johnnyseeds.com Kuehn , Michael Harris-Moran Seed Co, 25757 County Rd 21A, Esparto, CA, USA, 95627 m.kuehn@harrismoran.com Lewis, Mark Sakata Seed America, 105 Boronda Rd, Salinas, CA, USA, 93907 mlewis@sakata.com Liao, Charle Farmer Seed and Ag Co., Ltd., P.O. Box 45, Siu Swei, TAIWAN, 504 farmerseeds2000@yahoo.com.tw Liedl, Barbara WVSU, 201 ACEOP Admin Bldg, PO Box 1000, Institute, WV, USA 25112-1000 liedlbe@wvstateu.edu Majde, Mansour Gautier Semences, Route d' Avignon, Eyragues, FRANCE, 13630 mansour.majde@gautiersemences.com Maris, Paul DeRuiter Seeds, R&D NL BV, Leeuwenhoekweg 52, Bergschenhoek, THE NETHERLANDS, 2661CZ carla.schoonus@deruiterseeds.com Massoudi, Mark Ag Biotech Inc., P.O. Box 1325, San Juan Bautista, CA, USA,

95045 info@agbiotech.net Maxwell, Douglas P. University of WI, Madison, 7711 Midtown Rd, Verona, WI,

USA, 53593 douglas.maxwell08@gmail.com

104

McCaslin, Mark FLF Tomatoes, 18591 Mushtown Rd, Prior Lake, MN, USA, 55372 mccaslin@integra.net McGlasson, Barry University of Western Sydney, Centre for Plant and Food Science, Locked Bag 1797, Penrith South DC, NSW, AUSTRALIA, 1797 b.mcglasson@uws.edu.au McGuire, Cate Arcadia Biosciences, Inc. , 220 Cousteau Pl Ste #105, Davis, CA, USA, 95618 Merk, Heather Penn State University., Dept of Horticulture, 103 Tyson Building, University Park, PA, USA, 16802 hlml192@psu.edu Min, Chai Beijing Vegetable Research Center (BVRC), PO Box 2443, Beijing, PEOPLES REPUBLIC of CHINA, 100089 chaimin@nercv.com Myers, Jim Oregon State University, Dept. of Horticulture, rm 4017, Ag & Life Sci

Bldg., Corvallis, OR, USA, 97331 myersja@hort.oregonstate.edu Nadal, Michael Danson Seed Co, 10851 Woodbine St, Los Angeles, CA, USA, 90034-7675 Nakamura, Kosuke Kagome Co. Ltd., 17 Nishitomiyama, Nasushiobarashi,

Tochigi, JAPAN, 329-2762 Kosuke_Nakamura@kagome.co.jp North Carolina State University, NCSU Library, Campus Box 7111, Raleigh, NC, USA,

27695-0001 Ouyang, Wei Magnum Seeds, Inc., 5825 Sievers Road, Dixon, CA, USA, 95620 weiouyang1@yahoo.com Ozminkowski , Richard Heinz N.A., PO Box 57, Stockton, CA, USA, 95201 Rich.Ozminkowski@us.hjheinz.com Panthee, Dilip R. N.C. State U., Mountain Hort Crops Res & Ext Center,

455 Research Dr, Mills River, NC, USA, 28759 dilip_panthee@ncsu.edu Peters , Susan Nunhems USA, 7087 E. Peltier Rd., Acampo, CA, USA, 95220 susan.peters@nunhems.com Picard, Madame Florence Vilmorin, Service documentation, Route du Manoir,

La Menitre, FRANCE, 49250 Purdue University Library TSS, Unit Serials, 504 W. State St, West Lafayette, IN, USA,

47907-2058

105

Randhawa, Parm California Seed and Plant Lab, 7877 Pleasant Grove Rd, Elverta, CA, USA, 95626 randhawa@calspl.com Rascle, Christine Clause Tezier, Domaine de Maninet, Route de Beaumont, Valence, FRANCE, 26000 christine.rascle@clausetezier.com Saito, Atsushi National Institute of Vegetable and Tea Science, 360 Kusawa, Ano, Tsu, JAPAN, 514-2392 ashin@affrc.go.jp Sasaki, Seiko Plant Breeding Station of Kaneko Seeds, 50-12,

Furuichi-machi 1-chome, Maebashi City,Gunma, JAPAN, 371-0844 Scott, Jay University of Florida, Gulf Coast Research and Education Center, 14625

County Rd 672, Wimuama, FL, USA, 33598 jwsc@ufl.edu Semences, Gautier Gautier Semences, BP1, 13630, EYRAGUES, FRANCE recherche@gautiergraines.fr Semillas Fito c/ Selva de Mar 111, Barcelona, SPAIN, 8019 info@semillasfito.com Seno, Akiyoshi American Takii Inc., 11492 S. Ave D, Yuma, AZ, USA, 85365 aseno@takii.com Sharma, R.P. University of Hyderabad, Dept. of Plant Sciences, School of Life

Sciences, Hyderabad, INDIA, 500 046 rpssl@uohyd.ernet.in Shintaku, Yurie 2-10-2, Shimizu, Suginami-ku, Tokyo, JAPAN, 167-0033 Shupert, David Syngenta Seeds, 10290 Greenway Rd, Naples, FL, USA, 34114 david.shupert@syngenta.com Stack, Stephen Colorado State University., Biology, 1878 Campus Delivery, Fort

Collins, CO, USA, 80523-1878 sstack@lamar.colostate.edu Stamova, Liliana 1632 Santa Rosa St., Davis, CA, USA, 95616 listamova@yahoo.com Stamova, Boryana 2825 Bidwell St, Apt 4, Davis, CA, USA, 95618 Stevens, Mikel Brigham Young University, 275 Widtsoe Bldg, PO. Box 25183,

Provo, UT, USA, 84602 mikel_stevens@byu.edu Stoeva-Popova, Pravda Winthrop University, Department of Biology, 202 Life

Sciences Building, Rock Hill, SC, USA, 29732 stoevap@winthrop.edu

106

Stommel, John USDA-ARS, Genetic Improvement Fruits & Vegetables Laboratory, Bldg. 010A, BARC-West, 10300 Baltimore Ave., Beltsville, MD, USA, 20705 john.stommel@ars.usda.gov Takizawa, Kimiko Japan Horticultural Production and Research Inst., 2-5-1 Kamishiki Matsudo-shi, Chiba, JAPAN, 270-2221 takizawa@enken.or.jp Thomas, Paul 4 Juniper Court, Woodland, CA, USA, 95695 Thome, Catherine United Genetics Seeds Co., 764 Carr Ave., Aromas, CA, USA, 65004 cathy@unitedgenetics.com Tong, Nankui Campbell's Soup Co, Veg. Research & Dev Center, 28605 County

Road 104, Davis, CA, USA, 95691 Nankui_Tong@Campbellsoup.com University of California Riverside, Serv/Serials Technical, PO Box 5900, Riverside, CA,

USA, 92517-5900 University of New Hampshire Library, Serials Unit, 18 Library Way, Durham, NH, USA, 03824-3520 University of Wisconsin, Steenbock Library, 550 Babcock Dr, Madison, WI, USA, 53706 mquigley@library.wisc.edu van Schriek, Marco Keygene N.V., P.O. Box 216, Wageningen,

THE NETHERLANDS, 6700AE marco.van_schriek@keygene.com

Vecchio, Franco Nunhems Italy SRL, Via Ghiarone 2, Sant' Agata, Bolognese (BO), ITALY, 40019 Verbakel, Henk Nunhems Netherlands BV, R& D Library, PO Box 4005, Haelen, THE NETHERLANDS, 6080 AA S.vanRoy@nunhems.com Vinals, Fernando Nuez COMAV, Ciudad Politecnica de la Innovacion, Edificio 8-E.,

Excalera J. 3a Planta, Camino de Vera S/N, Valencia, SPAIN, 46022 fnuez@btc.upv.es

Volin, Ray Western Seed Americas, Inc., 15165 Dulzura Ct, Rancho Murieta, CA, USA, 95683-9120 ray@westernseed.com Wang, Wendy Xi'an Jinpeng seeds co. ltd., A803 of YuDao Hua Cheng,

No. 8 Feng Cheng 1 road, Xi'an City, Shaan'xi, PR CHINA, 710018 MASTER@JINPENGSEED.COM WA State University Libraries, SEA Serial record, 100 Diary Rd, Pullman, WA, USA 99164-0001

107

AUTHOR INDEX Chetelat, R. T. 66

Daunay M.C. 6

Fulladolsa, Ana Cristina 41

García, Brenda E. 41

Gilberston, R. L. 54

Hanson P 54

Laterrot H. 54

Lofty, Christopher 58

Maxwell, Douglas P. 41, 54

Mejía, Luis 41, 54

Melgar, Sergio 41

Méndez, Luis 41,54

Rivera, V.V. 54

Sánchez, Amilcar 54

Scott, J.W. 6

Secor, G.A. 54

Smith, Julian 58

Stoeva-Popova, Pravda 58

Teni, Rudy E. 41

Wang J.-F 6