GENETIC CONSIDERATIONS IN ECOSYSTEM RESTORATION USING NATIVE TREE SPECIES .••, The development of forest tree seed zones in the Pacific Northwest of the United States Brad St Clair United States Department of Agriculture Forest Service, Pacific Northwest Research Station, United States Seed zones and seed movement guidelines con­ tribute to the restoration of native ecosystems by ensuring adapted and resilient plant popula­ tions. Seed zones have a long history in the Pacific Northwest of the United States. Plantation for­ estry was initiated in the early twentieth century with the establishment of the Wind River Nursery by the United States Forest Service in 1910. The nursery was establ ished in southwestern Wash­ ington State to reforest and restore large areas of bare land and understocked forests result­ ing from large forest fires and logging. Initially, foresters did not pay particular attention to the off-site sources increased as stands aged, with a particularly sharp increase in the years after an extreme cold-weather event. These observations led in the 1940s to the es­ tabl ishment of the first seed collection zon es and seed collection gu idelines for Douglas-fir. A sys­ tem to cert ify the sta nd orig in of forest tree seed was established by the mid-1960s, and in 1966 seed zone maps for Washington and Oregon were published. These maps were widely used and have served their purpose of ensuring adapted planting stock for reforestation and restoration. In the meantime, rese archers have learned much source of forest tree seed. Seed came from eas­ ily accessible locations, often lower elevation more about geographic patterns of genetic vari­ ation in adaptive traits for a va riety of forest tree species, primaril y from short-term genecological studies such as those by Campbell {1986) and by forests near population centres and at logging operations. By the 1930s, however, it was be­ coming evident that not all plantations were as productive as they could be, particularly those at higher elevations, when compared with adja­ cent natura ll y regenerated stands. The gradual Sorensen (1992). Genecological studies co nsider genetic variation as found in common garden trials, and relate that variation to the cl imates or physiography of seed sources . Consistent, sensi­ decline of trees from non-local sources was also ble correlations between genetic variation and evident in two pioneering research studies begun in 1912: the Wind River Arboretum, wh ich tested seed-source environments indicate that a trait has responded to natural selection and ma y be of trees from around the world for their suitability io the Pacific Northwest, and the Douglas-Fir He­ redity Study, which addressed questions of type and location of Douglas-fir {Pseudotsuga men­ ziesii (Mirb.) Franco) parents from which to collect seed . Mortality and poor growth of trees from adaptive importance. Based on results from gene­ colog ical studies, se ed zones in Washington and Oregon were revised, primarily enlarging them in latitudinal directions, but mostly maintaining ele­ vation lim its (Randall, 1996; Randall and Berrang, 2002). This is because forest trees in temperate 49 THE STATE OF THE WORlD'S FOREST GENETIC RESOURCES- THEMAT IC STU DY and boreal regions are pr im arily adapted to mini­ mum winter temperatures, which are largely as­ maintaining genetic d iversity and structure of for­ est trees at landscape scales that are likely most sociated with elevation. Adaptation to aridity is also important in some regions, and may be par­ ticularly important in the trop ics. Species may also d iffer in the scale and patterns of genetic va ria­ tion and adaptation, and the revised seed zones in Oregon and Washington take t hese differences into account. Some species, such as Douglas-fir, are tightly adapted to their environments and may be considered specialist species . Other spe­ cies, like western red cedar (Thuja plicata Donn ex D. Don), are more generally adapted and may be considered genera li st species. Consequently, Douglas-fir has many seed zo nes with relatively narrow elevation bands of 150 m, whereas west­ ern red cedar has fewer seed zones with elevation bands of 450 to 600 m (Figure 15-1). An additional benefit of seed zones is that they contribute to important for adaptation. Seed zones and seed -movement guidelines have he lped to ensure productive, healthy and di­ ve rse fo rests in the Pacific Northwest for the past half-century and mo re. What have we learned? First, even with limited or no knowledge of genetic structure of a species, a reasonable as­ sumption is that native populations are at least approximately adapted to their local environ­ ments (Savolainen, Pyhajarvi and KnOrr, 2007) . The question then becomes, how local is local? Start somewhere. Make assumptions about the climatic or other environmental variables that are most important for adaptation , and deline­ ate seed zones based on those assumptions. In the Pacific Northwest. initial seed zones were based on climatic variables of cold and drought, Figure 15-1. Seed zones for Douglas-fir and western red cedar in Oregon and Wash in gton State, United States for Douglas-fir Source: adapted from Randall (1996); Randall and Berrang (2002). 50 western red cedar GENETIC CO NSI DERATIONS IN ECOSYSTEM RESTORATION USING NAT IVE TREE SPECI ES vegetation t ypes and p hysiography, especially el­ References evation . Genecolog y studies indicated th at many of those zones were too conservative, part icularly in t he north- south direction and particularly for some species that were later determined to be generalists. Revised seed zones t ook int o account t his new knowledge, but in t he meantime, or igi­ nal se ed zon es based primaril y on cl imat e served Cam pbell, R.K. 1986. Mapped genetic vanat1on of Douglas-fir to guide seed transfer 1n southwest Oregon . Silvae Genet., 35: 85-96 . Randall, W.K. 1996 . Forest tree seed zones for western Orego n . Salem, OK, USA, Oregon Department of Forestry. their purpose. Second, short-t erm g en ecology studies are val­ uable for indicating genetic structure important f or adaptati on and for delineating seed zones Randall, W.K. & Berrang, P. 2002. Washin g ton tree seed tran sfer zones. Olympia, WA, USA, Washington Department of Natural Resou rces. and seed-movement guidelines. These may be f ollowed by longer-t erm reciprocal transplant studies or provenance tests to evaluate long­ term adaptive respon ses, including estimat ­ Savolainen, 0 ., Pyhajarvi, T. & Knurr, T. 2007. Gene flow and local adaptation in trees. Annu. Rev. Ecol. Evo/. Syst. , 38: 595-619. ing productivity g iven climates at the locations of seed sources and plant ing sites (see Wang, O'Neill and A itken, 2010) . An important f inding from these studies is that each species must be cons idered individuall y, and that t he pattern s and scale of adaptation are not always obvious beforehand. Finally, during t he last fe w decades, scientists and land manag ers have recogn ized t hat cl i­ mates are changin g and have begun to conside r management responses. Knowledge of genet ic variation in adapt ive traits is important for un­ derstanding responses of n at ive population s t o Sorensen, F.C. 1992 . Genetic varia tion and seed transfer guidelines for lodgepole pine m cen tral Oregon . Research Paper PNW-RP-468. Portland, OR, USA, USDA Forest Service, Pacific Northwest Resea rch Station. Wang, T., O'N eill, G.A. & Aitken, S.N. 2010. Integrating environmental and genetic effects to predict responses of trees to clima te. Ecol. Appl., 20: 153-163 . climate change and f or evaluating managem ent option s to adapt t o climate change, includin g planting populations adapted to future climates and ensuring genetic diversity for future evolu­ t ion. The primary lesson from the development of seed zones in the Pacific Northwest is that, rather t han w aiting for the genetic knowledge t o accu­ mulate, it is bett er to act based on the best avail­ able knowledge, w hich may be from other spec ies in other regions, and th en to adjust management responses based on new knowledge from genetic studies. 51 Recommended citation: Bozza no, M., Jalonen. R.. Thomas. E., Boshier. D., Gallo, L., Cavers, S., Bordiics, S., Smith, P. & Loo, J., eds. 2014. Genetic considerations in ecosystem restoration using native tree species. State of the World's Forest Genetic Resources- Thematic Study. Rome, FAO and Bioversity International. Photo credits: p. 47 A. 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