APS_Jan2023

Volume 77

JANUARY 2023

Number 1

AMERICAN POMOLOGICAL SOCIETY F ounded in 1848 I ncorporated in 1887 in M assachusetts

2022-2023

PRESIDENT K. GASIC

FIRST VICE PRESIDENT P. CONNER

SECOND VICE PRESIDENT M. OLMSTEAD

TREASURER A. ATUCHA

EDITOR R. P. MARINI

SECRETARY L. DEVETTER

RESIDENT AGENT MASSACHUSETTS W. R. AUTIO

EXECUTIVE BOARD

N. BASSIL Past President

K. GASIC President

P. CONNER 1 st Vice President

M. OLMSTEAD 2 nd Vice President

L. DEVETTER Secretary

TOM KON ('19 - '23)

GINA FERNANDEZ ('21 - '24)

DAVID KARP ('22 - '25)

ADVISORY COMMITTEE 2020-2023 B. BYERS M. DOSSETT A. PLOTTO E. VINSON D. WARD 2021-2024 J. SAMTANI D. TRINKA S. MEHLENBACHER M. FARCUH G. BRAR 2022-2025 C. LUBY M. MUEHLBAUER L. REINHOLD A. WALLIS S. YAO

CHAIRS OF STANDING COMMITTEES

Editorial R. PERKINS-VEAZIE Wilder Medal Awards B. BLACK

Shepard Award F. TAKEDA Nominations M. PRITTS

Membership M. PRITTS

U. P. Hedrick Award E. FALLAHI

Website M. OLMSTEAD

Registration of New Fruit and Nut Cultivars J. PREECE, K. GASIC, D. KARP

1

January 2023

Volume 77 CONTENTS

Number 1

Published by THE AMERICAN POMOLOGICAL SOCIETY Journal of the American Pomological Society (ISSN 1527-3741) is published by the American Pomological Society as an annual volume of 4 issues, in January, April, July and October. Membership in the Society includes a volume of the Journal. Most back issues are available at various rates. Paid renewals not received in the office of the Business Manager by January 1 will be temporarily suspended until payment is received. For current membership rates, please consult the Business Manager. Editorial Office: Manuscripts and correspondence concerning editorial matters should be addressed to the Editor: Richard Marini, 203 Tyson Building, Department of Plant Science, University Park, PA 16802-4200 USA; Email: richmarini1@gmail.com. Manuscripts submitted for publication in Journal of the American Pomological Society are accepted after recommendation of at least two editorial reviewers. Guidelines for manuscript preparation are the same as those outlined in the style manual published by the American Society for Horticultural Science for HortScience, found at http://c.ymcdn.com/sites/www.ashs.org/resource/resmgr/files/style_manual.pdf . Postmaster: Send accepted changes to the Business office. Business Office : Correspondence regarding subscriptions, advertising, back issues, and Society membership should be addressed to the Business Office, C/O Heather Hilko, ASHS, 1018 Duke St., Alexandria, VA 22314; Tel 703-836 4606; Email: ashs@ashs.org Page Charges : A charge of $50.00 per page for members and $65.00 per page ($32.00 per half page) will be made to authors. In addition to the page charge, there will be a charge of $40.00 per page for tables, figures and photographs. Society Affairs : Matters relating to the general operation of the society, awards, committee activities, and meetings should be addressed to Michele Warmund, 1-31 Agriculture Building, Division of Plant Sciences, University of Missouri, Columbia MO 65211; Email:warmundm@missouri.edu. Society Web Site : http://americanpomological.org Diversity of Pawpaw (Asimina triloba) Cultivars in USDA Repositories and Selected Retail Nurseries c. 2022 – Richard B. Frost. ................................................................................................. 2 Performance of ‘Modi ® ’ apple trees on several Geneva rootstocks managed organically: Five-year results from the 2015 NC-140 Organic Apple Rootstock Trial – Terence Bradshaw, Wesley Autio, Suzanne Blatt, Jon Clements, Todd Einhorn, Rachel Elkins, Esmail Fallahi, Poliana Francescatto, Jaume Lordan, Ioannis Minas, Gregory Peck, Terence Robinson, and Shengrui Yao...................... 14 Genetic Analysis of Resistance to Pythium ultimum a Major Component of Replant Disease in Apple Rootstocks – Gennaro Fazio, Mark Mazzola, and Yanmin Zhu............................................. 28 About the Cover: ‘Brewster’ lychee.................................................................................................. 38 Maurice Adin Blake: Father of the New Jersey Peach Industry – Richard P. Marini. ...................... 39 Low-Temperature Survival of 'Creshaven' Peach Flower Buds and Fruit Yield on Eight Rootstocks in the 2017 NC-140 Rootstock Trial – Michele R. Warmund, Mark Ellersieck, James R. Schupp . .... 51 Index for Volume 76 .......................................................................................................................... 62 Instructions to Authors. ..................................................................................................................... 64

J ournal of the A merican P omological S ociety

2

Journal of the American Pomological Society 77(1): 2-13 2023

Diversity of Pawpaw (Asimina triloba) Cultivars in USDA Repositories and Selected Retail Nurseries c. 2022 R ichard B. F rost Additional index words: ancestry, fruit weight, genomic-morphologic associations Abstract This study reviewed available data for Pawpaw ( Asimina triloba ) cultivars in the U.S. with the goal of determin ing genetic diversity and genomic-morphologic relationships. Ninety cultivars available at U.S. retail sites and 20 cultivars at USDA sites (6 unique from retail) in 2022 are listed. Nine genetic studies of pawpaw are then reviewed, finding 17 genetic associations among the retail cultivars. Recent agricultural, chemical, tensile, and spectrometric morphology data are also reviewed. An association with larger fruit weight and cultivars derived from ‘Middletown’ was found, but the remaining forms of morphology data were either uncorrelated with genom ic data or otherwise unsuitable to determine genomic associations. A discussion of criteria for future genomic morphologic studies of Asimina triloba is also included.

Over the last 140 years the development and preservation of U.S. Pawpaw ( Asimina triloba ) selections has passed through many hands, including J.A. Little, E.J. Downing, G..A. Zimmerman, O.E. White, G.L. Slate, C. Davis, J. Gordon, P. Thomson, R.N. Pe terson, J. Lehman, and C. England (Davis, 1982; England, 2022; Little, 1905; Peterson, 1991; Peterson, 2003; Pomper et al., 2009; Thomson, 1974; Zimmerman, 1941). A few unusual cultivars are now appearing in the retail trade, including variegated ‘Spilt Milk’ plus three freestone cultivars ‘Cantaloupe’, ‘Honey Dew’, and ‘Marshmallow’. Much has been written about the fruit in the last few decades, including food chemistry (Brannan et al., 2015; Grygorieva et al., 2021), propa gation and planting guides (Cothron, 2021; Hummer, 2020; Tabacu et al., 2020), and ecotours (Moore, 2015). Interest in Pawpaw cultivation has also spread overseas (Bran nan and Coyle, 2021). A list of cultivars cur rently held at USDA repositories is given in Table 1 and those presently available from U.S. retail nurseries in Table 2. This report is part of a series involving

genetic ID and genetic clades within lesser studied fruits. In tandem it has been discov ered that unsound mathematical practices have found their way into mainstream bioin formatics and are currently considered valid by investigators and reviewers alike (Frost, 2022b). Several of the articles reviewed here are no exception. The primary issues en countered here are use of non-metric dissimi larities, pair-grouping, and use of data with missing values for distance. Note that “met ric” here refers to the mathematical definition of distance – not units of measure. All three of these practices appear in biology curricula and consequently the authors cited here have used them unwittingly. Materials and Methods Genomic studies. Nine genomics studies of U.S. Pawpaw cultivar diversity have been conducted since 1990. Three were authored by H. Huang et al: the first primarily to de termine an advanced set of RAPD markers (Huang et al., 2000), the second utilizing 71 RAPD markers on 37 cultivars (Huang et al., 2003), and the third using ALFP markers

1 Frost Concepts, Vista CAUSA. email: tangent.vectors@gmail.com orcid: https://orcid.org/0000-0002-0214-4230

s

P awpaw

3

List of cultivars currently held at USDA repositories (USDA, 2022).

Table 1. List of cultivars currently held at USDA repositories (USDA, 2022).

Cultivar Allegheny Ames 3129 Ames 32220 Ames 7465

Year introduced USDA Repository Administration

2007 1984 2014 1986 1990 1990 1970 2014 ?

NCGR Corvallis

National Arboretum

North Central Regional PI Station North Central Regional PI Station

Anniston

NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis

KSU Atwood KSU Benson

Mango

NA 82138 NA 83885

National Arboretum National Arboretum

?

NC-1

1976 1950

NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis NCGR Corvallis

Overleese

PA Golden No. ? 1986

Potomac Prolific

1994 1985 1990 1990 1968 1994

Shenandoah Susquehanna

Taytwo Wabash

(Wang et al., 2005). Unfortunately, all three applied a non-metric measure to determine dissimilarities. Of these the second publica tion contains all marker data but also suf fered from missing marker values. A recent mathematics study has salvaged 45 from the set (Table 3), providing a coarse measure of genomic dissimilarity among the specimens (Frost, 2022a). A second set of published studies are affili ated with Kentucky State University. The first study (Pomper et al., 2003) used question able marker loci which under Jaccard’s met ric produces 4 sets of zero distances between cultivars known to have different parentage. The second (Pomper et al., 2010) is plagued by poor data quality and use of a non-metric dissimilarity measure. A comparison of ge netic associations and clades found by Frost (2022a) and the second KSU study is shown in Table 4. The third study (Lu et al., 2011) used 20 SSR primer pairs but unfortunately did not publish their marker data. A fourth study (Botkins et al., 2012) used 6 standard

7 An interesting undergraduate study from West Virginia University analyzed 19 un named specimens from 3 campus pawpaw patches for clonal variation using 12 ISSR microsatellite markers (Fontana, 2019). A heat map was used to visualize the inter-patch diversity, shown here in a topological graph (Figure 1). In 2021 a trio from the University of Georgia published a study of morphologic and microsatellite data from 20 U.S. sites as sociated with pre-Columbian settlements and 62 possibly wild specimen sites in the eastern U.S. (Wyatt et al., 2021). Unfortunately the investigators chose non-metric dissimilarity measures to analyze their data and the mark er values contain numerous missing values (Table 5) making them unsuitable for any analysis (Schlueter and Harris, 2006). Morphology studies. Several studies of pawpaw cultivar morphology data have been published in the past 20 years. Survival rates SSR loci to estimate clonal variation in 7 lo calized wild or feral pawpaw patches but also did not publish the marker data.

4 Table 2. Cultivars available at some point in 2022 from one or more of the following U.S. retail sites: Cricket Hill Garden, Elmore Roots Nursery, England's Orchard, Hidden Springs Nursery, Just Fruits and Exotics, Kiefer Nursery NC, Nash Nurseries, One Green World, Peaceful Heritage Nursery, Perfect Circle Farm, Raintree Nursery, Red Fern Farm, Restoring Eden, Tollgate Gardens. J ournal of the A merican P omological S ociety Table 2. Cultivars available at some point in 2022 from one or more of the following U.S. retail sites: Cricket Hill Garden, Elmore Roots Nursery, England's Orchard, Hidden Springs Nursery, Just Fruits and Exotics, Kiefer Nursery NC, Nash Nurseries, One Green World, Peaceful Heritage Nursery, Perfect Circle Farm, Raintree Nursery, Red Fern Farm, Restoring Eden, Tollgate Gardens. Table 2. Cultivars available at some point in 2022 from one or more of the following U.S. retail sites: Cricket Hill Garden, Elmore Roots Nursery, England’s Orchard, Hidden Springs Nursery, Just Fruits and Exotics, Kiefer Nu s ry NC, Nash Nurseries, One Gree World Peaceful He itage Nursery, Perfect Circle Farm, Raintree Nursery, Red Fern Farm, Restoring Eden, Tollgate Gardens. 8-20 9-58-? Al Horn White Flesh Allegheny Asterion Atria

8-20 Avatar

9-58-? Belle

Al Horn White Flesh Benny's Favorite Benny's Favorite Cluster

Allegheny Betria

Asterion Canopus Canopus Convis

Atria Cantaloupe Cantaloupe Davis Gainsville #2 Davis Gainsville #2 IXL

Avatar Carmelo Dr. Chill Carmelo Dr. Chill Gatria

Belle Caspian

Betria Collins

Caspian Ford Amend Ford Amend Golden Moon Golden Moon Jerry's Delight Jerry's Delight Lady D Lady D Marshmallow Marshmallow Overleese Prima 1216 Overleese Prima 1216 Rigel Spilt Milk Variegated Susquehanna Rigel Spilt Milk Variegated Taylor Susquehanna

Cluster Free Byrd

Collins Fulbright's Delight

Convis Gainsville #1 Gainsville #1 Honey Dew Honey Dew KSU Benson

Free Byrd Greenriver Belle Greenriver Belle Kentucky Champion

Fulbright's Delight Halvin KSU Atwood Halvin

Gatria Jerry's Big Girl Jerry's Big Girl LA Native

KSU Chappell

IXL

KSU Atwood Lehman's Chiffon Lehman's Delight Lynn's Favorite KSU Benson

KSU Chappell Mango

Kentucky Champion Mitchell

LA Native Maria's Joy

MSU Golden

NC-1

Mango Nyomi's Delicious

Lehman's Chiffon Lehman's Delight Lynn's Favorite

Maria's Joy October Moon October Moon Potomac

PA Golden No. ? PA Golden No. 1 PA Golden No. 2 PA Golden No. 3

Mitchell Prolific

MSU Golden Quaker Delight SAA Zimmerman-#? Summer Delight Quaker Delight SAA Zimmerman-#? Summer Delight Sweet Alice

NC-1

Nyomi's Delicious Rebecca's Gold

Rappahannock

PA Golden No. ? PA Golden No. 1 PA Golden No. 2 PA Golden No. 3

Potomac Regulus

Prolific SAA-Overleese SAA-Overleese Sue

Rappahannock Shenandoah

Rebecca's Gold Sibley

Regulus Sidewinder Sidewinder Sunsprout Sunsprout Tallahatchie Tallahatchie VE-21

Shenandoah Sunflower

Sibley Sunglo

Sue Sweet

Sunflower Sweet Potato Sweet Potato Tropical Treat Tropical Treat Windstar

Sunglo Sweet Virginia Sweet Virginia UVM #1

Sweet Taytwo Walters Taytwo

Sweet Alice Tollgate

Taylor Wabash

Tollgate Wells

UVM #1 Zimmerman

VE-21

Wabash

Walters

Wells

Windstar

Zimmerman

Table 3. Names of 45 RAPD marker primer sequences used in coupled analysis of pawpaw genetic markers and ancestry records (Frost, 2022a). Table 3. Names of 45 RAPD marker primer sequences used in coupled analysis of pawpaw genetic mark ers and ancestry records (Frost, 2022a). Table 3. Names of 45 RAPD marker primer sequences used in coupled analysis of pawpaw genetic m rk rs and ancestry records (Frost, 2022a). A07-1600 A07-0600 A11-0850 A11-0600 A11-0425 A12-0550 B07-1200 B07-0550 B08-0900 B09-0900 B10-1775 B10-1200 B10-0950 B10-0900 B11-0525 C02-0650 C04-1300 C04-1675 C 4 175 C 8 425 C 15 C 3 13 C 5 1050 C 5 6 D 5 5 D 5 0 D 5 45 D 5 6 D05 05 D 5 055 D 5 425 D 6 10 D 6 D16 40 D16 0 25 D20 07 D20 00 E 1 850 E0 04 E 1 675 E 4 08 E 70 E16 05 E16 1025 E17 90 A07-1600 A07-0600 A11-0850 A11-0600 A11-0425 A12-0550 B07-1200 B07-0550 B08-0900 B09-0900 B10-1775 B10-1200 B10-0950 B10-0900 B11-0525 C02-0650 C04-1300 C04-1675 C04-1175 C08-0425 C11-1550 C13-1300 C15-1050 C15-0650 D05-1250 D05-0500 D05-0450 D05-0600 D05-0575 D15-0550 D15-0425 D16-1000 D16-0525 D16-0400 D16-0325 D20-0775 D20-1100 E01-0850 E01-0450 E11-1675 E14-0850 E15-0700 E16-0550 E16-1025 E17-0900

of specimens in irrigated and unirrigated orchards were recorded by investigators at Kentucky State University (Pomper et al., 2008). Of those in the irrigated plot for which there is also viable genomic data, only ‘Over leese’, ‘Susquehanna’, ‘Taylor’, and ‘Wells’

8 had survival rates in the range 75% to 88% and the remainder had no casualties. Aver age fruit weights at harvest were recorded by investigators at KSU, Ohio University, and North Carolina Cooperative Extension (Fig ure 2). The KSU investigators also contribut- 8

Figures

P awpaw

5

Figure 1. Genetic diversity among 19 numbered specimens from 3 separate pawpaw patches studied by K. Fontana (Fontana, 2019). Dissimilarity values computed by a metric in package GenAlEx (Peakall and Smouse, 2018) and plotted here in a least bridges graph (Frost, 2022b). Circle markers are from patch 1, squares are from patch 2, and triangles from patch 3. Spatial orientation and scale in the graph are arbitrary. Figure 1. Genetic diversity among 19 numbered specimens from 3 separate pawpaw patches studied by K. Fontana (Fontana, 2019). Dissimilarity values computed by a metric in package GenAlEx (Peakall and Smouse, 2018) nd plotted here in a least bridges graph (Frost, 2022b). Circle markers ar from patch 1, squares are from patch 2, and triangles from patch 3. Spatial orientation and scale in the graph are arbitrary.

ed ag iculturally important Growing Degree Days data (Pomper et al., 2008) (see Table 6). Measurements of interest in food science were taken in Ohio (Brannan et al., 2015) of 4 applicable cultivars (see Table 7). Results Genetic associations. Seventeen genetic associations determined from a reduced set of 49 RAPD markers (Frost, 2022a) are listed in column 3 of Table 4. However, the speci mens in this study are a biased selection of Asimina triloba genomes and so the smallest association groups defined here should be tak en with a grain of salt. More study is clearly needed to determine better methods of genetic discrimination among Pawpaw cultivars. Genomic-morphologic connections. Pos sible connections between genomic and

12 Hopes for finding a relationship between the Growing Degree Days (GDD) data and genomic groups were not met. Linear and multilinear models constructed from the KSU clades and genetic associations of this study returned correlation coefficients of -0.05, 0.09, and 0.098. The disparity between the marker groups and GDD data is illustrat ed in Figure 3. Success occurred with comparisons of ge nomic data and vetted fruit weight data. The raw data originated in 4 separate studies as morphologic data were examined for this study. The small variances in survival rates were considered problematic for analysis. The phenolic and spectrometric data were also not analyzed due to caution in the study paper regarding different levels of ripeness among the specimens.

6 J ournal of the A merican P omological S ociety Table 4. Comparison of genetic associations and clades determined by R. Frost a (Frost, 2022a) and investigators at KSU b (Pomper et al., 2010). The determination of genetic associations is illustrated in Figure 5. Pivots were selected for their independence and utility with other cultivars. Some of the groups are due to unique ancestors while others (e.g. 3-11) either share an unknown ancestor or a common set of chromosomal subsequences that have been reinforced by breeding programs. Table 4. Comparison of genetic associations and clades determined by R. Frost a (Frost, 2022a) and inves tigators at KSU b (Pomper et al., 2010). The determination of genetic associations is illustrated in Figure 5. Pivots were selected for their independence and utility with other cultivars. Some of the groups are due to unique ancestors while others (e.g. 3-11) either share an unknown ancestor or a common set of chromo somal subsequences that have been reinforced by breeding programs.

Year introduced

In 2022 retail trade

Selected genomic pivots a

Genomic associations a

Primary association a

KSU clade b

Cultivar

Middletown

1915 1985 1990 1990 1990 1990 1990 1994 1994 1950 1994 1994 1990 1976 1994 1945 1994 1985 1985 1968 1968 1980s 1970 1994 1994 1994 1974 1990 1990 1979 1986 1990 1994 1990 1990 1990

X

A A A A A A A A A

A A A A A A A A A A A A A A A B B B B C C C D D D D E E E

IV

Prolific

R

II

2-49 9-47

untested

III III

Rappahannock Shenandoah Susquehanna

R R R

V II

5-5

IV IV

7-90

Overleese Potomac

R R

ADF ADF ADG AEF

V

III IV

3-21 2-54

II V II

NC-1

R

AF AG

2-10

Sweet Alice

R

X

B B

III

9-58-2

untested untested untested

SAA-Zimmerman-1 SAA-Zimmerman-2

BA

BDF

Taylor Taytwo Wilson

R R

X

C C C D D D E E E F F F G

I

V

I

Sunflower

R

X

V V

1-68

Wabash

R R R

III

8-20

DB

II V

Rebecca's Gold

X

9-58-1

untested untested

Wells-PPF

Mitchell

R R R

X

F F F

V

PA-Golden

untested

Wells

IV

3-11

X

G G

II II

11-13 10-35

GA

numerous numerous

n/a n/a

III

1-23

V

9

7 P awpaw Table 5. Percent missing values across sites, specimens, and markers in a population dispersal study at University of GA (Wyatt et al., 2021). Table 5. Percent missing values across sites, specimens, and markers in a population dispersal study at University of GA (Wyatt et al., 2021). Table 5. Percent missing values across sites, specimens, and markers in a population dispersal study at University of GA (Wyatt et al., 2021). Site Population Types Sites mi sing values Specime s issing values Marker allele missing values Table 5. Percent missing values across sites, specimens, and markers in a population dispersal study at University of GA (Wyatt et al., 2021). Site Population Types Sites missing value Specimens missing value Marker alleles missing values Anthropomorphic 70.0% 52.8% 61.1% Wild Table 6. Measurements of GDD by KSU investigators (Pomper et al., 2008). Some of the estimated peak flowering weeks have been back-calculated from GDD and harvest week. Table 6. Measurements of GDD by KSU investigators (Pomper et al., 2008). Some of the estimated peak lowering weeks have been back-calculated from GDD and harvest week. Table 6. Measurements of GDD by KSU investigators (Pomper et al., 2008). Some of the estimated peak flowering weeks have been back-calculated from GDD and harvest week. Table 6. Measurements of GDD by KSU investigat r ( omper et al., 2008). Some of th estimated peak flowering weeks have been back-calculated from GDD and harvest week. Cultiv rs in retail circulation Estimate peak flowering week at KSU sites GDD at KSU sites Peak harvest week at KSU sites Site Population Types Sites missing values Anthropomorphic 70.0% Wild 82.3% Wild 82.3 Specimens missing values 52.8% 70.1% 70.1 Marker alleles missing values 61.1% 72.2% 72.2 Anthropomorphic 70.0% 82.3% 52.8% 70.1% 61.1% 72.2%

Cultivars in retail circulation Cultivars in retail circulation PA-Golden Wabash Rappahannock PA-Golden Wabash Rappahannock NC-1 PA-Golden Overleese Wabash NC-1 Rappahannock Overleese Taytwo Taylor Taytwo Shenandoah NC-1 Overleese Taylor Shenandoah Susquehanna Susqu hanna Pot mac

Estimated peak flowering week at KSU sites Estimated peak flowering week at KSU sites 16 16 16 16 16

GDD at KSU sites GDD at KSU sites 2499 2572 2499 2572 2586

Peak harvest week at KSU sites Peak harvest week at KSU sites 36 36 36 36 36

16 16 16 16 16 16 16 16 16 15 15

2499 2572 2586 2620 2637 2648 2676 2697 2703 2720 2736 2737 2751 2753 720 2586 2620 2637 2648 2676 2697 2703 2720 2736 2737 2751 2753 2620 2637 2648 2676 2697 2703 2736 2737 2751 2753

37 37 37 37 37 37 37 37 37 37 37

16 16 16 16 16 16 16 16 16 16 15 15

36 37 37 37 37 37 37 37 37 37 37 37

16 16 16 16 16 16 16 16 16 16 16 16 15 15

36 36 36 37 37 37 37 37 37 37 37 37 37 37

Taytwo Taylor Shenandoah Susquehanna Potomac Mitchell Sunflower Mitchell Sunflower Wells Potomac Mitchell Sunflower Wells 8-20 8-20

Wells Table 7. Soluble s lids concentrations, phenolic, tensile, and spectrometric measurements of pawpaw cultivars (Brannan et al., 2015) applicable to H. Huang’s RAPD markers (Huang et al., 2003). Table 7. Soluble solids concentrations, phenolic, tensile, and spectrometric measurements of pawpaw cul tivars (Brannan et al., 2015) a licable to H. Huang’s RAPD markers (Huang et al., 2003). 8-20 Table 7. Soluble solids concentrations, phenolic, tensile, and spectrometric measurements of pawpaw cultivars (Brannan et al., 2015) applicable to H. Huang’s RAPD markers (Huang et al., 2003). Table 7. Soluble solids concentrations, phenolic, tensile, and spectrometric measurements of pawpaw cultivars (Brannan et al., 2015) applicable to H. Huang’s RAPD markers (Huang et al., 2003). C ltivar NC-1 25.7 5.68 ± 0.41 0.248 {62.9,-4.8,30.2} {77.1,10.1,45.9} pulp texture (kg) pulp texture (kg) skin CIE color (L*,a*,b*) skin CIE color (L*,a*,b*) pulp CIE color (L*,a*,b*) pulp CIE color (L*,a*,b*) Cultivar Soluble solids conc. Soluble solids conc. 25.1 Phenolics (μmol/g) Phenolics (μmol/g) Overleese

Phenolics (μmol/g) 5.38 ± 0.67 0.415 6.21 ± 0.20 0.363 5.38 ± 0.67 0.415 6.21 ± 0.20 .363 5.68 ± 0.41 0.248 5.30 ± 0.17 0.198 5.38 ± 0.67 0.415 6.21 ± 0.20 0.363 5.68 ± 0.41 0.248 5.30 ± 0.17 0.198 5.30 ± 0.17 0.198

skin CIE color (L*,a*,b*) {63.3,-7.8,35.2} {75.1,6.7,42.0} {61.7,-6.0,29.6} {71.2,12.2,53.2} 3.3,-7. ,35.2} {75.1,6.7,42.0} 1.7,-6.0,29.6} {71.2,1 .2,53.2 pulp CIE color (L*,a*,b*) {62.9,-4.8,30.2} {77.1,10.1,45.9} {65.1,-8.8,33.0} {79.3,2.1,34.6} {63.3,-7.8,35.2} {75.1,6.7,42.0} {61.7,-6.0,29.6} {71.2,12.2,53.2} wise comparison, a common ordinal speci men was selected, e.g. ‘Overleese’ compared to the Brannan and Greenawalt series. The percentage of weight change of each speci men from the ordinal was then calculated for {62.9,-4.8,30.2} {77.1,10.1,45.9} {65.1,-8.8,33.0} {79.3,2.1,34.6} {65.1,-8.8,33.0} {79.3,2.1,34.6}

NC-1

25.7

Rebecca's Gold

23.5

Overleese Taytwo

Soluble solids conc. 25.1 23.5 25.2

pulp texture (kg)

Rebecca's Gold

Cultivar

Taytwo

25.2

NC-1

25.7 25.1 23.5 25.2

Overleese Rebecca's Gold shown in Figure 2. The data were filtrated by making pairwise comparisons between the Greenawalt series and the others – under the assumption that fruit weights from different sites vary by linear proportion. For each pair Taytwo

10

10

J ournal of the A merican P omological S ociety

8

Figure 2. Fruit production data in average grams per fruit from Oxford NC, SW OH, and KSU sites in KY sorted by L. Greenawalt's SW OH data (Brannan et al., 2015; Cantaluppi and Coley, 2020; Greenawalt et al., 2019; Pomper et al., 2008). Figure 2. Fruit production data in average grams per fruit from Oxford NC, SW OH, and KSU sites in KY sorted by L. Greenawalt’s SW OH data (Brannan et al., 2015; Cantaluppi and Coley, 2020; Greenawalt et al., 2019; Pomper et al., 2008). Figure 2. Fruit production data in average grams per fruit from Oxford NC, SW OH, and KSU sites in KY sorted by L. Greenawalt's SW OH data (Brannan et al., 2015; Cantaluppi and Coley, 2020; Greenawalt et al., 2019; Pomper et al., 2008).

Figure 3. Disparity between GDD data and genetic marker groups. Figure 3. Disparity between GDD data and genetic marker groups.

each of the two series, e.g. the % gains for the Brannan series using ‘Overleese’ as or dinal are {-30%,0%,~10.8%,~56.9%}. The differences of percentage gain series were examined and any specimen pair with more than ±10% difference was rejected. For ex ample, the series differences between Bran nan and Greenawalt were approximately

{-5.6%,0.%,6.4%,49.4%} and thus specimen ‘NC-1’ was eliminated from that pair of se ries. Any specimens for which there was no comparative value was also eliminated. Fig ure 4 illustrates the vetted results from all 4 studies. The vetted data was used to construct the genomic-morphologic comparison of Ta ble 8. The influence of cultivar ‘Middletown’ 13

Figure 3. Disparity between GDD data and genetic marker groups.

13

P awpaw

9

Figure 4. Vetted fruit weight (g) data extracted from Figure 2. Figure 4. Vetted fruit weight (g) data extracted from Figure 2.

Figure 5. Illustration of genetic association selection for cultivar 8-20. Only genetic distances less than the sub-average distance of 12 mismatches were considered (see Figure 6). Spatial orientation and line seg ment lengths likely have no correlation to actual 45-dimensional space. Figure 5. Illustration of genetic association selection for cultivar 8-20. Only genetic distances less than the sub-average distance of 12 mismatches were considered (see Figure 6). Spatial orientation and line segment lengths likely have no correlation to actual 45-dimensional space.

on higher fruit weights is apparent. The culti var ‘Taylor’ and its possible sibling ‘Taytwo’ both appear in section of lower weights, as do ‘Sunflower’ and ‘Mitchell’. These latter two are also present as minor components in

14 the higher fruit weight specimens. One can speculate here that the influence of ‘Middle town’ is too dominant for them but a future study with more refined data is probably war ranted.

ment lengths likely have no correlation to actual 45-dimensional space.

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ure 6. Sample distribution of marker mismatches ! in 36 unique specimens measured with 45 error RAPD markers of H. Huang (Frost, 2022a). Figure 6. Sample distribution of marker mismatches in 36 unique specimens measured with 45 error-free RAPD markers of H. Huang (Frost, 2022a).

Figure 7. Topological graph of genomic associations among 36 Pawpaw cultivars of this study. Solid lines are nearest neighbor connections. Dashed lines are cultivar associations of genomic pivots determined by least bridges graph (Frost, 2022b). Line lengths and spatial orientation are arbitrary. Figure 7. Topological graph of genomic associations among 36 Pawpaw cultivars of this study. Solid lines are nearest neighbor connections. Dashed lines are cultivar associations of genomic pivots determined by least bridges graph (Frost, 2022b). Line lengths and spatial orientation are arbitrary.

11 Table 8. Comparison of vetted average fruit weights with genomic groupings by R. Frost a (above) and KSU b (Pomper et al., 2008) Table 8. Comparison of vetted average fruit weights with genomic groupings by R. Frost a (above) and KSU b (Pomper et al., 2008) P awpaw

Fruit weights in SW Ohio (g)

Year introduced

Genomic associations a

KSU clade b

Cultivar Mitchell Taylor Taytwo

117 119 121 148 153 160 167 172 194

1979 1968 1968 1970 1990 1950

F

V

C C D A

I

V V V V V V II

Sunflower Shenandoah

Overleese

ADF

Rebecca's Gold 1974

E

NC-1

1976 1990

AF

Susquehanna

A

Discussion The specimens in the genomic studies of H. Huang (Huang et al., 2000; Huang et al., 2003) are for the most part closely related due to a century of breeding programs. This situation produces a condition of “too much cohesion” in topological graphs using met ric distances (Frost, 2022b). As such, these graphs are difficult if not impossible to par tition with standard graph theory methods. The approach taken here of genomic pivots (pseudo basis points) is one alternative (see Table 4 and Figure 5). However, the asso ciations alone do not provide an adequate “map” of specimen relations. Figure 7 shows an attempt to resolve the issue with a hybrid graph, incorporating associations with near est neighbor relations. If the USDA online records are correct then the USDA germplasm repositories for Asimina triloba poorly represent genomic diversity in the species. One would expect specimens representing each of the genomic pivots identified above plus others selected for traits of agricultural interest. Viable ge netic fingerprinting of the USDA collection would be beneficial. The application of 45 markers from H. Huang’s original set (Huang et al., 2003) to fruit weights show that they have merit be yond ancestral relations. Using the entire set of 71 on all cultivars in retail circulation

could provide a more exacting view of diver sity within the selections and guidance for future breeding. If the fingerprinting is to be effective, the RAPD data for each specimen needs to be composed of one error-free set or 5-8 sets with 10% or less missing values and enough overlap to produce a high-confidence correlated error-free set (Frost, 2022b). If an investment is made in taking new genomic measurements, it would be highly beneficial to collect an array of morphologic data in-situ. Ripe fruit for laboratory assay should be obtained from each of the leaf specimen trees and some quantitative mea sure of “ripeness” should be made for reg istration of compound concentrations in the fruit samples (Brannan et al., 2015). Com pounds of interest in the fruit include annon acins, carbohydrates, fruit sugars, flavonoids, glutamates, phenols, and proteins. Tensile tests should include skin shear strength and bulk texture. Average seed counts and per cent by volume are desirable for selective breeding. Collection of harvest degree-days information (fruit set date, harvest date, tree location) and cultivar vigor would be a bonus. The testing of annonacin concentra tions is important for understanding possible health risks of the fruit. A determination can be made by comparing annonacin concentra tions to lifetime dosage limits for injectable annonacin used in contact treatment of can

11

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cer cells times the expected percent of drug escape into the patient blood stream. Conclusions Categorizing cultivar traits with genomic groupings in Pawpaw is difficult with cur rently available information. It has been demonstrated here that prior studies using RAPD analysis can only provide coarse group distinctions and that nearly all prior genomic studies are based on invalid math ematical approaches – albeit no fault of the authors. A re-analysis of Pawpaw genomics and morphological characteristics over many cultivars (100+) is certainly in order. Hope fully the current revolution in long-read se quencing technology will provide cost-effec tive means of analysis in the future. Acknowledgements All computations and analyses were per formed with Wolfram Mathematica® v.13. The author also expresses his gratitude to the editor and reviewers for their helpful guid ance. Literature Cited Botkins, J., K.W. Pomper, J.D. Lowe, and S.B. Crabtree. 2012. Pawpaw Patch Genetic Diversity, and Clonality, and its Impact on the Establishment of Invasive Species in the Forest Understory. Journal of the Kentucky Academy of Science 73:113-121. doi: https://doi.org/10.3101/1098-7096-73.2.113. Brannan, R.G. and M.N. Coyle. 2021. Worldwide Introduction of North American Pawpaw (Asimina triloba): Evidence Based on Scientific Reports. Sustainable Agriculture Research 10:1-19. doi: https://doi.org/10.5539/sar.v10n3p19. Brannan, R.G., T. Peters, and S.T. Talcott. 2015. Phytochemical analysis of ten varieties of pawpaw (Asimina triloba [L.] Dunal) fruit pulp. Food Chemistry 168:656-661. doi: https://doi. org/10.1016/j.foodchem.2014.07.018. Cantaluppi, C. and J. Coley. 2020. Pawpaw Cultivar Evaluation: 2007-2019. Journal of the NACAA 13. url: https://www.nacaa.com/journal/bba6a701 ede1-4eda-b557-c047c282b412. Cothron, B. 2021. Pawpaws: The Complete Growing and Marketing Guide. New Society Publishers. url: https://www.google.com/books/edition/Pawpaws/- tonEAAAQBAJ.

Davis, C. 1982. The Pawpaw in southern Michigan, p. 38-41. In: P. Thomson (ed.). California Rare Fruit Growers Yearbook 1982. CRFG, Bonsall CA. url: https://www.google.com/books/edition/Yearbook/ D-44AQAAIAAJ. England, C. 2022. England’s Orchard. http://www. nuttrees.net/. Fontana, K. 2019. Role of geographic distance in clonal propagation of Asimina triloba subpopulations of northern West Virginia. West Virginia University. url: https://researchrepository.wvu.edu/core_ arboretum_ug/3/. Frost, R. 2022a. Coupled analysis of Pawpaw (Asimina triloba) genetic markers and ancestry records. International Journal on Computational Science & Applications 12:1-8. url: https://frostconcepts.org/ articles/CoupledAnalysisAsiminaTriloba2022.pdf. Frost, R. 2022b. Decades of miscomputation in genomic clades and distances. International Journal on Computational Science & Applications 12. url: http://wireilla.com/papers/ijcsa/ V12N4/12422ijcsa01.pdf. Greenawalt, L., R. Powell, J. Simon, and R.G. Brannan. 2019. A retrospective analysis of pawpaw (Asimina triloba [L.] Dunal) production data from 2005-2012. Journal of the American Pomological Society 73:2-11. url: http://www.pubhort.org/ aps/73/v73_n1_a1.htm. Grygorieva, O., S. Klymenko, O. Vergun, K. Fatrcová Šramková, O. Shelepova, Y. Vinogradova, V.H. Sedláčková, and J. Brindza. 2021. Studies of the chemical composition of fruits and seeds of pawpaw (Asimina triloba (L.) Dunal). Agrobiodiversity for Improving Nutrition, Health and Life Quality 5. url: https://agrobiodiversity.uniag.sk/scientificpapers/ article/view/342/395. Huang, H., D.R. Layne, and T.L. Kubisiak. 2000. RAPD inheritance and diversity in pawpaw (Asimina triloba). Journal of the American Society for Horticultural Science 125:454-459. doi: https:// doi.org/10.21273/JASHS.125.4.454. Huang, H., D.R. Layne, and T.L. Kubisiak. 2003. Molecular characterization of cultivated pawpaw (Asimina triloba) using RAPD markers. Journal of the American Society for Horticultural Science 128:85-93. doi: https://doi.org/10.21273/ JASHS.128.1.0085. Hummer, K. 2020. NCGR Corvallis - Asimina Germplasm. USDA ARS. https://www.ars.usda. gov/pacific-west-area/corvallis-or/national- clonal germplasm-repository/docs/ncgr-corvallis-asimina germplasm/. Little, J.A. 1905. The Pawpaw. OG Swindler. url: https:// www.google.com/books/edition/The_Pawpaw_ Asimina_Triloba_a_Native_Frui/uJ0aAAAAYAAJ.

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Lu, L., K.W. Pomper, J.D. Lowe, and S.B. Crabtree. 2011. Genetic variation in pawpaw cultivars using microsatellite analysis. Journal of the American Society for Horticultural Science 136:415-421. doi: https://doi.org/10.21273/JASHS.136.6.415. Moore, A. 2015. Pawpaw: In Search of America’s Forgotten Fruit. Chelsea Green Publishing. url: https://www.google.com/books/edition/Pawpaw/ f3BOCgAAQBAJ. Peakall, R. and P.E. Smouse. 2018. GenAlEx website. Australian National University. https://biology assets.anu.edu.au/GenAlEx/Welcome.html. Peterson, R.N. 1991. Pawpaw (Asimina). Genetic Resources of Temperate Fruit and Nut Crops 290:569-602. url: https://www.actahort.org/ books/290/290_13.htm. Peterson, R.N. 2003. Pawpaw variety development: a history and future prospects. HortTechnology 13:449-454. doi: https://doi.org/10.21273/ HORTTECH.13.3.0449. Pomper, K., S. Crabtree, J. Lowe, and J. Lehman. 2009. Pawpaw and the American Persimmon in Workshop 13 - Native Fruits of the Midwest. ASHS 106th Annual Conference St. Louis MO. url: https://journals.ashs.org/hortsci/view/journals/ hortsci/44/4/article-p919.xml. Pomper, K.W., S.B. Crabtree, S.P. Brown, S.C. Jones, T.M. Bonney, and D.R. Layne. 2003. Assessment of genetic diversity of pawpaw (Asimina triloba) cultivars with intersimple sequence repeat markers. Journal of the American Society for Horticultural Science 128:521-525. doi: https://doi.org/10.21273/ JASHS.128.4.0521. Pomper, K.W., S.B. Crabtree, D.R. Layne, R.N. Peterson, J. Masabni, and D. Wolfe. 2008. The Kentucky Pawpaw Regional Variety Trail. Journal American Pomological Society 62:58. url: https:// www.pubhort.org/aps/62/v62_n2_a3.htm. Pomper, K.W., J.D. Lowe, L. Lu, S.B. Crabtree, S. Dutta, K. Schneider, and J. Tidwell. 2010.

Characterization and identification of pawpaw cultivars and advanced selections by simple sequence repeat markers. Journal of the American Society for Horticultural Science 135:143-149. doi: https://doi.org/10.21273/JASHS.135.2.143. Schlueter, P.M. and S.A. Harris. 2006. Analysis of multilocus fingerprinting data sets containing missing data. Molecular Ecology Notes 6:569 572. doi: https://doi.org/10.1111/j.1471 8286.2006.01225.x. Tabacu, A.F., A.C. Butcaru, J. Lehman, and F. Stănică. 2020. Top grafting response of some pawpaw (Asimina triloba Dunal) genotypes. Scientific Papers. Series B, Horticulture:198-203. url: http:// horticulturejournal.usamv.ro/pdf/2020/issue_1/ Art31.pdf. Thomson, P. 1974. California Rare Fruit Growers Yearbook, vol. 6. CRFG, Bonsall CA. url: https:// frostconcepts.org/references/CRFG1974Yearbook. pdf. USDA. 2022. Asimina triloba - GRIN Global. USDA ARS. https://npgsweb.ars-grin.gov/gringlobal/ taxon/taxonomydetail?id=4485. Wang, Y., G.L. Reighard, D.R. Layne, A.G. Abbott, and H. Huang. 2005. Inheritance of AFLP markers and their use for genetic diversity analysis in wild and domesticated pawpaw [Asimina triloba (L.) Dunal]. Journal of the American Society for Horticultural Science 130:561-568. doi: https://doi. org/10.21273/JASHS.130.4.561. Wyatt, G.E., J. Hamrick, and D.W. Trapnell. 2021. The role of anthropogenic dispersal in shaping the distribution and genetic composition of a widespread North American tree species. Ecology and evolution 11:11515-11532. doi: https://doi. org/10.1002/ece3.7944. Zimmerman, G. 1941. Hybrids of the American papaw. Journal of Heredity 32:83-93. doi: https:// doi.org/10.1093/oxfordjournals.jhered.a105006.

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Journal of the American Pomological Society 77(1): 14-27 2023 Performance of ‘Modi ® ’ apple trees on several Geneva rootstocks managed organically: Five-year results from the 2015 NC-140 Organic Apple Rootstock Trial T erence B radshaw 1 , W esley A utio , S uzanne B latt , J on C lements , T odd E inhorn , R achel E lkins , E smaeil F allahi , P oliana F rancescatto , J aume L ordan , I oannis M inas , G regory P eck , T erence R obinson , A nd S hengrui Y ao Additional index words: yield efficiency, tree survival, trunk cross-sectional area, tree size, cumu lative yield, crop load, fruit weight Abstract In 2015, an orchard trial of ten apple rootstocks was established at ten locations in the United States and Canada using ‘Modi®’ as the scion cultivar. Trees were managed in accordance with United States organic standards to expose these rootstocks to the nutrient conditions and biome typically associated with organic tree-fruit produc tion. Rootstocks included nine named Cornell-Geneva clones [Geneva® 11 (G.11), Geneva® 30 (G.30), Ge neva® 41 (G.41), Geneva® 202 (G.202), Geneva® 214 (G.214), Geneva® 222 (G.222), Geneva® 890 (G.890), Geneva® 935 (G.935), and Geneva® 969 (G.969)] and M.9 NAKBT337. All trees were spaced 1 x 3.5 m and trained using the tall spindle system. After 5 years, the greatest mortality was for trees on M.9 NAKBT337 (14%). Rootstocks separated into size classes from large semi-dwarf to small dwarf. G.890 resulted in large semi-dwarf trees, and G.202 produced moderate semi-dwarfs. G.41 and G.30 resulted in small semi-dwarf trees, and trees on G.935 were large dwarfs. G.11, G.214, G.969 and M.9 NAKBT337 resulted in trees that were moderate dwarfs, and G.222 resulted in small dwarf trees. The most yield efficient (cumulatively, 2016-19) trees in the trial were on G.935, G.11, and G.969, and the least efficient trees were on G.202 and G.890. The largest fruit (2016-19) were harvested from trees on G.30, G.41, G.890, and M.9 NAKBT33, and the smallest were harvested from trees on G.202.

NC-140 is a Multi-state Research Project organized by state agricultural experiment stations, USDA, and agencies in Mexico and Canada. During the 45 years of its ex istence, it has evaluated nearly all new tem perate, tree-fruit rootstocks utilizing uniform trials in diverse locations in North America (Cowgill et al., 2017). All prior NC-140 trials were managed with “conventional”, i.e., non organic, programs. This nomenclature of “or ganic” vs. “conventional’ suggests that there are standard management practices applied across each system, when in reality, there are a large number of practices and intensi ties of management that are applied within and across systems. Because of this potential variation in management, in this paper we re

fer to “organic” compared to “non-organic” systems, as conventions may vary among each system across regions and cooperators. In 2019, 947 million pounds (~430,000 mt) of certified organic apples were produced in the U.S., with about 93% of that production located in Washington state (NASS, 2020). While relatively small-scaled compared to overall apple orchard land use, organic pro duction still constitutes substantial commer cial activity in diverse apple producing states throughout the U.S., and organic orchards may be found in all apple-producing states. Organic production presents several po tential limitations to overall orchard perfor mance. In the U.S., organic apple production is more common in western states that have

1 Corresponding author: Terence L. Bradshaw, University of Vermont, Plant and Soil Science, Horticulture Re search and Education Center, 63 Carrigan Dr, Burlington, VT 05405, tbradsha@uvm.edu

A pple

15

less summer precipitation than in the more humid mid-west and eastern U.S. A trial in California comparing organic to convention al production found few differences between the systems, and although trees were slightly smaller, and profitability was greater for the organically-managed orchard (Swezey et al., 1998). In Washington, apple yields under organic management were generally compa rable with non-organic apples in multiyear studies (Peck et al., 2006; Reganold et al., 2001), which explains the relatively greater proportion of organically-grown fruit in that state than in most other U.S. states or Cana dian provinces. Because of increased pre cipitation and humidity that leads to greater disease pressure and more insect pest species present, organic apple production is substan tially more difficult in the eastern U.S. than in drier western states. In a long-term evalu ation of organic apple production in Ver mont, cumulative crop yield was far below economically acceptable conventional yields for nearly all cultivars trialed, and newly es tablished trees were all unprofitable (Brad shaw et al., 2016a; Bradshaw et al., 2016b). In Kentucky, yield in a long-term trial of or ganically-managed, scab-resistant apple cul tivars (SRCs) was substantially lower than what is typical for non-organic, commercial apples in the state, and only 43-64% of fruit were considered marketable primarily due to insect and disease damage (Williams et al., 2015). Evaluation of organic apple produc tion in New York has shown greater success. In one study, the SRC ‘Liberty’ was evaluat ed over four seasons in comparative organic and integrated (a hybrid of organic and con ventional) management systems and overall, organic management was competitive with integrated fruit production for yield and tree growth, although pest incidence was gener ally greater (Peck et al., 2010). In a separate trial that compared two intensities of organic management with a non-treated control in an orchard consisting of multiple SRCs, similar levels of pest and disease incidence to the prior New York and Vermont studies were

observed (Agnello et al., 2017). However, higher prices received for certified-organic fruit would likely offset a lower percentage of clean fruit under organic management, so long as yield is sufficient as outlined by Bradshaw et. al. (2016b). Over the last several years, several new rootstocks have been released from the Cor nell-Geneva breeding program (managed jointly by Cornell University and the United States Department of Agriculture-Agricul tural Research Service). Many of these Ge neva series rootstocks have been previously evaluated in other NC-140 trials (Autio et al., 2013; Autio et al., 2017a; Autio et al., 2020a; Autio et al., 2017b; Autio et al., 2020b; Autio et al., 2011b; Autio et al., 2011c; Marini et al., 2014; Robinson et al., 2007). The objec tives of this current trial were to assess and compare performance of several Cornell Geneva rootstocks managed using organic management procedures at multiple sites in North America. Materials and Methods In spring, 2015, an orchard trial of 10 apple rootstocks was established at 10 sites in North America (Table 1) under the coor dination of the NC-140 Multi-State Research Committee. ‘Modi®’ [a U.S. trademark of ‘CIVG198’(Leis et al., 2008)] was used as the scion cultivar, and trees were propagat ed by Wafler Nursery (Wolcott, NY, USA). ‘Modi®’ is a ‘Gala’ x ‘Liberty’ hybrid SRC bred in Italy. This cultivar was selected for its reported high fruit quality, consistent yield, and resistance to fire blight. Rootstocks in cluded nine named Cornell-Geneva clones [Geneva® 11 (G.11), Geneva® 30 (G.30), Geneva® 41 (G.41), Geneva® 202 (G.202), Geneva® 214 (G.214), Geneva® 222 (G.222), Geneva® 890 (G.890), Geneva® 935 (G.935), and Geneva® 969 (G.969)] and M.9 NAKBT337. The trial was planted in California, Colo rado, Idaho, Massachusetts, Michigan, New Mexico, Nova Scotia (Canada), New York (Geneva and Ithaca), and Vermont. In eight