Identification of Historic Apple Trees in the Southwestern United States and Implications for Conservation
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Hop i Sill yii 4400 2009. tamed apple culttvars introduced to or selected in North America. Identification of Historic Apple Trees in Modern commercial apple production requires consistency of ripening time, quality retention during processing and shipping, and the Southwestern United States and long storage life, and not all varieties can meet these criteria (Golanij and Bauer. 2004). Implications for Conservation Market pressures have reduced the diversity Kanin J. Routson' of fnut trees once grown in small fiumily orchards- -where diversity of ripening time, Arid Lands Resource Sciences, Unii'ei'sit-t' ofA rizona, 1955 East Si.itli Street, sizes, textures, and flavors were celebra- P.O. Box 210184, Tucson, AZ 85719 ted—to only a few handfuls of commonly planted commercial cultivars. Curi'enil y. I Ann A. Reillev, Adam D. Henk, and Gayle M. Volk apple cultivars account for over 90% of the National ('enic,' fin' Genetic Resources Preservation, U. S. Depai'/rnenr of apples sold in the United States, with 'Red Agriculiui'e, Fort Collins, CO 80521 Delicious' constituting 41% of this hgure Dennis, 2008). In The Fruit. Bern' and A'te! Ai/i/ioonal iiidrv words. .tioliv. simple sequence repeat. genetic diversity J;ii anton- ( Whcalv. 2001) Kent Whealy lists 1500 apple varieties cun'entiv available Abstract. lany apple varieties commonl y planted in the United States a century ago can through U.S. nurseries, many of which have no longer be found in toda y 's orchards and nurseries. Abandoned farmsteads and been developed through modern fruit breed- historic orchards harbor considerable agrobiodiversity , but the extent and location of ing. This suggests a substantial decrease in that (li%ersitv is poorl y understood. We assessed the genetic diversit y of 280 apple (Ma/xis the number of apple cultivars offored through xdo,,iestiea Borkh.) trees growing in 43 historic f'ai'nisktad and orchard sites in Arizona, U.S. nurseries over the past century (779/a by Utah, and New Mexico using seven microsatellite markers. We compared the samples to Ragan's calculations and 89% by Bussey's), 109 cultivars likel y introduced into the southwest in the late 1901 and earl y 20th although we do not know to what extent this centuries. Genetic anal y sis revealed 144 genoty pes represented in the 280 field samples. naming actuall y represented genetically dis- \\e identified 34 of these 144 genot y pes as cultivars brought to the region bN , Stark tinct cultivars. Brothers Nursery and b y USDA agricultural experiment stations. One hundred twenl y of Although tue loss of'on-farni diversity can the total samples (43%) had DNA fingerprints that suggested the y were representative of be lasting, fruit trees have all over these 34 ctiltivars. The remaining 160 samples—representing 110 genotspes—had annual crops because these trees can live to unique fingerprints that did not match all the fingerprinted cultivars. The results remarkably old ages, surviving some fads in of this study confirm for the first time that a high diversit y of historic apple genotypes consumer demand. Sin gle apple trees have remain in homestead orchards in the U.S. southwest. Future efforts targeting orchards in been known to live 150 years or longer, in the southwest should focus on conservation for all unique genot y pes as a means to sustain triany areas, it is still possible to find trees of both cultural heritage and biological genetic diversity. "IlCirloom" cultivars once abundant at the beginning of the 20th century. Remnant orchards planted before the "modern era" The late 19th century is often referred to desirable ciders. and those that were amena- of fruit production (Jackson. 2003) hang on as the golden years of apple growing in the ble to cooking and baking. This period of tenaciousl y around abandoned I'll rtii stead s United States (Calhoun. 1995: Hensle y, 2005). American horticultural history was preceded and historic orchards. Although farmstead Farmstead and kitchen orchards were planted by an era of fruit diversification, which trees often persist without their original with a wide variety of fruit trees to suit spanned much of the I 700s and into the early names being retained, they represent a snap- diverse family needs. Historically. rural live- I 800s. Thousands of trees emerged horn shot of the diversity of fruit varieties avail- lihoods were maintained b growing apples y seedling orchards, planted for cider and for able over a century ago during the peak of and pears that ripened in summer, would animal feed, that bore high-quality fruits fi'uit tree diversification. "keep" all winter in the cold cellar, produce worthy of naming. conserving, and distribut- Morphological and taxonomic traits typ- ing. These trees were clonally propagated ically used to differentiate between apple from cuttings and traded and sold to become eultivars call ambiguous as a result of Received for publication 16 Dec. 2008. Accepted elements of the diverse orchards of the 19th the broad phenotypic variation under differ- for publication 6 Feb. 2009. century (Beach. 1905: Hedrick, 1950). The ent environmental influences. Furthermore, This project was pat of Kanin Routson's Master's last remnants of 19th century plantings may many 19th century apple cultivars are mor- research at Northern Arizona Universit y. Thank still he alive in landscapes and tire phologically similar to one another and accu- you to Gary Nahhan, Thomas Sisk-, David Osterg- the subject of this regional southwestern rate descriptions are often lacking. making ren, Gery Allan, and other staff at NAU who helped survey. with this study. We also express our gratitude to all conventional identification methods difficult, who have provided funding for this project: the USDA pomologist W.H. Ragan under- if not impossible, for these centur y-old trees. National Park Service and the Margaret T. Morris took the task of recording the names and Genetic fitigerprinti ng. including micro- Foundation. [it we are grateful to Lee characteristics of every apple cultivar grown satellites, have become powerful and accu- Calhoun, Ram Fishman, and Philip Forsline for in the United States during the 19th century. rate tools for analyzing genetic diversity sending leaf samples of named varieties used in this In his book, The Nonzenc/ai,o-e q/ the Apple (Hammer et al., 2003). Microsatel lite loci, study and to Christopher Richards for use of (Ragan. 1905). Ragan lists 6654 unique or simple sequence repeats, are short nlicle- facilities at USDA-ARS-NCGRP. Lastly, we thank named apple varieties that lie found refer- otide sequences of up to six hasepairs Gerard Geenen of the Plant C yiornetry Services enced in U.S. literature between the years repeated in Schipnde I. The Netherlands) for determining ploidy tandem, head to tail, without 1804 and 1904. In 1980, [)an l3usse y began for the samples and reference culiivars. interruption. They are highly polymorphic. expanding on Ragan 's register to update Mention of trade nanles or coillmc'rcj at products in codominant markers that have been detected this article is solely for the purpose of providing descriptions of known cultivars and include in every organism thus far studied (I lancock, specific information and does not imply reeom- additional apple cultivars referenced in the 1999). Gianfranceschi et al. (1998), l-Iokanson niendation or endorsement by the U.S. Department U.S. literature up to the year 1980. Over a et al. (1998), and 1-Jemmat et al. (2003) were of Agriculture. decade after beginning this project, Bussey instrumental in developing Ma/us-specific To whom reprint requests should be addressed: (in press) is close to releasing The App/c' in microsatellites in the United States. In e-nlai I kjr53aemail .arizona.edu. North America, which lists over 14,000 Europe. Si lfverberg-Di lworth ci al. (2006) H0RTSClrNcE Vor. 44(3) JUN! 2009 589 and the II igli-Qual ity Disease Resistant Apples for Sustainable Agriculture have been forefront in developing microsatellites for Ma/us. In a similar study in Spain. Pereira- Lorenzo et al. (2007) evaluated the genetic diversity of 114 Spanish apple cultivars comparing the local Spanish landrace geno- types with 26 commercial apple cultivars found in the region. Similar to this study. Pereira-Lorenzo et al. (2007) found high Utah Colorado levels of genetic diversity in apple trees in I northern Spain based on observed heterozy- 42-43 • 41 gosity. Capitol Reef •" In this study, we sampled tissues from 280 apple trees from 43 historic sites within Arizona. Utah. and New Mexico to assess their diversity. Genotypes identified using seven highly variable markers were com- Taos 38-40 pared with reference genotypes of known 4 eultivars. The term "historic" is used in this text to refer to farmstead and orchard sites 26-33 Santa Fe planted during the late 19th and early 20th 25 Albuquerque centuries, and the term "heirloom" refers to Flagstaff cultivars introduced during the 19th century 21-24 .. and before as opposed to recent introductions Prescott 34-37 . developed through modern fruit breeding programs. 1-19 • 20 Materials and Methods Arizona I New Mexico Field co//eciwns. We collected leaf sam- ples from 280 apple trees located in 43 historic orchard sites on public and private (hut not tribal) lands in Arizona, New Mexico, and Utah (Fig. 1). Sampling took place from June through September of 2007. We targeted Fi g 1. I lisioric apple collection sites (numbered and noted h (tiamonds ) in Utah, Arizona. and Ncs places with presumed historic orchards and Mexico were sampled. Cities are provided for reference purposes (star designation). trees Of Visually differing morphologies. Local experts provided site location information. Sampling permission was obtained and sam- pies were used only for this study, not for gene num tags were nailed to the trees with a Polymerase chain reactions (P(.'Rs) were banking or crop improvement.