{{Short description|Pine tree found in western North America}} {{Redirect|Sugar Pine|the community in Madera County, California|Sugar Pine, California}} {{Speciesbox | name = Sugar pine | image = Sugar Pine.jpg | status = LC | status_system = IUCN3.1 | status_ref = <ref name="iucn status 13 November 2021">{{cite iucn |author=Farjon, A. |date=2013 |title=''Pinus lambertiana'' |volume=2013 |article-number=e.T42374A2976106 |doi=10.2305/IUCN.UK.2013-1.RLTS.T42374A2976106.en |access-date=13 November 2021}}</ref> | display_parents = 3 | parent = Pinus subsect. Strobus | genus = Pinus | species = lambertiana | authority = Douglas | range_map = Pinus lambertiana range map 1.png | range_map_caption = Natural range of ''Pinus lambertiana'' }}
'''''Pinus lambertiana''''' (commonly known as the '''sugar pine''') is the tallest and most massive pine tree and has among the longest cones of any conifer. It is native to coastal and inland mountain areas along the Pacific coast of North America, as far north as Oregon and as far south as Baja California in Mexico.
==Description==
=== Growth === The sugar pine is the tallest and largest ''Pinus'' species, commonly growing to {{convert|40|–|60|m|ft|round=5|sp=us}} tall, exceptionally to {{cvt|82|m|ft}} tall, with a trunk diameter of {{cvt|1.2|-|2.5|m|ftin}}, exceptionally {{cvt|3.5|m|ftin}}.<ref name="Arno">{{Cite book|last1=Arno|first1=Stephen F.|title=Northwest Trees: Identifying & Understanding the Region's Native Trees|last2=Hammerly|first2=Ramona P.|publisher=Mountaineers Books|year=2020|isbn=978-1-68051-329-5|edition=field guide|location=Seattle|pages=26, 30–35|language=en|oclc=1141235469|orig-date=1977}}</ref> The tallest recorded specimen is {{convert|83.45|m|ftin|abbr=on}} tall, is located in Yosemite National Park, and was discovered in 2015.<ref>{{cite news | title = 3 Sierra sugar pines added to list of 6 biggest in world | newspaper = Associated Press | location = South Lake Tahoe, California | language = English | date = 31 Jan 2021| url = https://apnews.com/article/nevada-forests-south-lake-tahoe-17501dc8cb094fc7ffb1543426631bc6 | access-date = 13 Feb 2023}}</ref> The second tallest recorded was "Yosemite Giant", an {{cvt|82.05|m|ftin}} tall specimen in Yosemite National Park, which died from a bark beetle attack in 2007. Yosemite National Park also has the third tallest, measured to {{cvt|80.5|m|ftin}} tall as of June 2013; the Rim Fire affected this specimen, but it survived. The next tallest known living specimens grow in southern Oregon; one in Umpqua National Forest is {{cvt|77.7|m|ftin}} tall and another in Siskiyou National Forest is {{cvt|77.2|m|ftin}} tall. [[File:Pinus lambertiana ProspectOR.jpg|thumb|left|upright|Old sugar pines in the Rogue River – Siskiyou National Forest, southern Oregon]]
The bark of ''Pinus lambertiana'' ranges from brown to purple in color and is {{Convert|5-10|cm|sp=us|frac=2}} thick.<ref name="Arno" /> The upper branches can reach out over {{Convert|8|m|abbr=on}}.<ref name="Arno" /> Like all members of the white pine group (''Pinus'' subgenus ''Strobus''), the leaves ("needles") grow in fascicles ("bundles") of five<ref name="Arno" /> with a deciduous sheath. They are {{cvt|5|–|11|cm|in|frac=4}} long.<ref>{{Jepson eFlora|38280|Pinus lambertiana}}</ref> Sugar pine is notable for having among the longest cones of any conifer, mostly {{cvt|10|–|50|cm|in|0}} long,<ref name="Arno" /><ref name="FNA">{{eFloras|1|233500939|Pinus lambertiana |family=Pinaceae |first=Robert |last=Kral}}</ref> although they can occasionally reach as much as {{cvt|56|cm|in|0}}.<ref name="u078">{{cite web | title=Oregon State University | website=Landscape Plants | url=https://landscapeplants.oregonstate.edu/plants/pinus-lambertiana | access-date=2025-09-20}}</ref><ref name="FEIS">{{cite web |last=Habeck |first=R. J. | title=Pinus lambertiana | website=US Forest Service | date=1990-10-26 | url=https://www.fs.usda.gov/database/feis/plants/tree/pinlam/all.html | access-date=2025-09-20}}</ref> Although the cones of the Coulter pine are more massive; the unripe weight of {{Convert|1-2|kg}} makes the sugar pine's cones perilous projectiles when chewed off by squirrels.<ref name="Arno" /> The seeds are {{cvt|1|–|2|cm|in|frac=4}} long, with a {{convert|2|–|3|cm|in|frac=4|adj=on|abbr=off|sp=us}} long wing<ref name="FNA" /> that aids their dispersal by wind. Sugar pine never grows in pure stands, always in a mixed forest, and is shade tolerant in its youth.<ref>{{Gymnosperm Database |family=Pinaceae |genus=Pinus |species=lambertiana}}</ref>
=== Distribution === The sugar pine occurs in Oregon and California in the Western United States southward to Baja California, specifically in the Cascade Range, Sierra Nevada, Coast Ranges, and Sierra San Pedro Martir. It is generally more abundant towards the south and can be found between {{Convert|500 and 1500|m|abbr=on}} above sea level.<ref name="Arno" />
===Genome=== The massive 31 gigabase mega-genome of sugar pine has been sequenced in 2016 by the large PineRefSeq consortium.<ref name=Stevens>{{cite journal|last=Stevens|first=K.A.|display-authors=et al.|date=2016|title=Sequence of the Sugar Pine Megagenome |journal=Genetics |volume=204|issue=4|pages=1613–1626|doi=10.1534/genetics.116.193227|pmid=27794028|pmc=5161289}}</ref> This makes the genome one of the largest sequenced and assembled so far.<ref name=Stevens/>
The transposable elements that make up the megagenome are linked to the evolutionary change of the sugar pine. The sugar pine contains extended regions of non-coding DNA, most of which is derived from transposable elements. The genome of the sugar pine represents one extreme in all plants, with a stable diploid genome that is expanded by the proliferation of transposable elements, in contrast to the frequent polyploidization events in angiosperms.<ref>{{Cite journal |last=Gonzalez-Ibeas |first=Daniel |display-authors=et. al|date=2016 |title=Assessing the Gene Content of the Megagenome: Sugar Pine (''Pinus lambertiana'') |journal=G3 (Bethesda) |volume=6 |issue=12 |pages=3787–3802|doi=10.1534/g3.116.032805 |pmid=27799338 |pmc=5144951 }}</ref>
thumb|right|Almost ripe female cones
=== Embryonal growth === In late stage of embryonal development, the sugar pine embryo changes from a smooth and narrow paraboloid to a less symmetric structure. This configuration is caused by a transverse orientation of division planes in the upper portion of the embryo axis. The root initial zone is established, and the epicotyl develops as an anlage flanked by regions of that define the cotyledonary buttresses. At this stage, the embryo is composed of the suspensor, root initials and root cap region, hypocotyl-shoot axis, and the epicotyl. The upper (distal) portion of the embryo, which gives rise to the cotyledons and the epicotyl, is considered to be the shoot apex.<ref>{{Cite journal |last=Berlyn |first=Graeme P |date=1967 |title=The Structure of Germination in Pinus Lambertiana Dougl. |journal=Yale School of Forestry & Environmental Studies, Bulletin Series |volume=77}}</ref>
=== Shoot apex === The shoot apex has the following four zones:<ref>{{Cite journal |last=Sacher |first=J.A. |date=1954 |title=Structure and Seasonal Activity of the Shoot Apices of Pinus Lambertiana and Pinus ponderosa. |journal=American Journal of Botany |volume=41 |issue=9 |pages=749–759|doi=10.1002/j.1537-2197.1954.tb14406.x |bibcode=1954AmJB...41..749S }}</ref> #The apical initials produce all cells of the shoot apex through cell division. It is located at the top of the meristem and the cells are larger in size compared to other cells on the surface layer. #The central mother cell generates the rib meristem and the inner layers of the peripheral tissue zone through cell division. It presents a typical gymnosperm appearance and is characterized by cell expansion and unusual mitosis that occurs in the central region. The rate of mitosis increases on its outer edge. #The peripheral tissue zone consists of two layers of cells that are characterized by dense cytoplasm and mitosis of high frequency. #Lastly, the rib meristem is a regular arrangement of vertical files of cells which mature into the pith of the axis.
==Etymology== Naturalist John Muir considered sugar pine to be the "king of the conifers". The common name comes from the sweet resin, which Native Americans used as a sweetener.<ref>{{cite web|url=http://www.oregonencyclopedia.org/articles/sugar_pine/#.VZr15PmqpBc|title=Sugar pine|website=Oregonencyclopedia.org|access-date=18 June 2017}}</ref> John Muir found it preferable to maple sugar.<ref name = Saunders>{{cite book |title=Edible and Useful Wild Plants of the United States and Canada |url=https://books.google.com/books?id=4IwrWyK7OmQC&q=Edible+and+Useful+Wild+Plants+of+the+United+States+and+Canada |last=Saunders |first=Charles Francis |year=1976 |publisher=Courier Dover Publications |isbn=0-486-23310-3 |page=219 }}</ref> It is also known as the great sugar pine. The scientific name was assigned by David Douglas in 1826,<ref name="Arno" /> in honor of the London botanist Aylmer Bourke Lambert.<ref>{{Cite book |last=Muir |first=John |title=The Mountains of California |date=2010 |publisher=Random House Publishing Group |others=Bill McKibben |isbn=978-0-375-75819-5 |series=Modern Library Classics Ser |location=Westminster |pages=112–114}}</ref>
==Ecology== === Wildlife === The large size and high nutritional value of the sugar pine seeds are appealing to many species. Yellow pine chipmunks (''Neotamias amoenus'') and Steller's jays (''Cyanocitta stelleri'') gather and hoard sugar pine seeds. Chipmunks gather wind-dispersed seeds from the ground and store them in large amounts. Jays collect seeds by pecking the cones with their beaks and catching the seeds as they fall out. Although wind is a main dispersal agent of sugar pine seeds, animals tend to collect and store them before the wind can blow them far.<ref>{{cite journal |last1=Thayer |first1=T |last2=Vander Wall |first2=S |year=2005 |title=Interactions between Steller's jays and yellow pine chipmunks over scatter-hoarded sugar pine seeds |journal=Journal of Animal Ecology |volume=74 |issue=2 |pages=365–374 |doi=10.1111/j.1365-2656.2005.00932.x |jstor=3505625 |doi-access=free |bibcode=2005JAnEc..74..365T }}</ref>
Black bears (''Ursus americanus'') feed on sugar pine seeds in the fall months within the Sierra Nevada. Both sugar pine and oak species are currently in decline, directly affecting black bear food sources within the Sierra Nevada.<ref>{{cite journal |last1=Mazur |first1=R |last2=Klimley |first2=AP |last3=Folger |first3=K |year=2013 |title=Implications of the variable availability of seasonal foods on the home ranges of black bears, ''Ursus americanus'', in the Sierra Nevada of California |journal=Animal Biotelemetry |volume=1 |issue=16 |page=16 |doi=10.1186/2050-3385-1-16 |doi-access=free|bibcode=2013AnBio...1...16M }}</ref>
=== Threats === The sugar pine has been severely affected by the white pine blister rust (''Cronartium ribicola''),<ref name="Moore2008">{{cite book |author1=Moore, Gerry |title=National Wildlife Federation Field Guide to Trees of North America |author2=Kershner, Bruce |author3=Craig Tufts |author4=Daniel Mathews |author5=Gil Nelson |author6=Spellenberg, Richard |author7=Thieret, John W. |author8=Terry Purinton |author9=Block, Andrew |publisher=Sterling |year=2008 |isbn=978-1-4027-3875-3 |location=New York |page=79}}</ref> a fungal pathogen accidentally introduced from Europe in 1909. A high proportion of sugar pines have been killed by the blister rust, particularly in the northern part of the species' range where blister rust has been present longer. The rust has also destroyed much of the Western white pine and whitebark pine throughout their ranges.<ref>{{cite web |last1=Maloney |first1=P |last2=Duriscoe |first2=D |last3=Smith |first3=D |last4=Burton |first4=D |last5=Davis |first5=D |last6=Pickett |first6=J |last7=Cousineau |first7=R |last8=Dunlap |first8=J |title=White Pine Blister Rust on High Elevation White Pines in California |url=http://www.caforestpestcouncil.org/2005%20CFPC%20Meeting/1506%20Maloney_Dunlap%2005BR%20Survey.pdf |archive-url=https://web.archive.org/web/20061009055647/http://www.caforestpestcouncil.org/2005%20CFPC%20Meeting/1506%20Maloney_Dunlap%2005BR%20Survey.pdf |archive-date=2006-10-09 |access-date=2007-02-05}}</ref> The U.S. Forest Service has a program for developing rust-resistant varieties of sugar pine and western white pine. Seedlings of these trees have been introduced into the wild. The Sugar Pine Foundation in the Lake Tahoe Basin has been successful in finding resistant sugar pine seed trees. Blister rust is much less common in California, where sugar, Western white and whitebark pines still survive in great numbers.<ref name="spf">{{cite web |title=Sugar Pine Foundation |url=http://www.sugarpinefoundation.org |access-date=18 June 2017 |website=Sugarpinefoundation.org}}</ref>
Sugar pine trees have been affected by the mountain pine beetle (''Dendroctonus ponderosae''), which is native to western North America. The beetles lay their eggs inside of the tree and inhibit the tree's ability to defend itself against invading species. Beetle infestation can also cause nutrient deficiencies that slowly weaken the tree's overall health, making pines more susceptible to other threats such as fires and white pine blister rust.<ref>{{cite web |year=2012 |title=Mountain pine beetle |url=http://www.invadingspecies.com/mountain-pine-beetle/ |url-status=live |work=Ontarios invading species awareness program |archive-url=https://web.archive.org/web/20200927011833/http://www.invadingspecies.com/mountain-pine-beetle/ |archive-date=2020-09-27}}</ref> Blister rust can weaken the tree and enable further infestation by mountain pine beetles.<ref>{{cite journal |last1=Van Mantgem |first1=PJ |last2=Stephenson |first2=NL |last3=Keifer |first3=M |last4=Keeley |first4=J |year=2004 |title=Effects of an introduced pathogen and fire exclusion on the demography of sugar pine |journal=Ecological Applications |volume=14 |issue=5 |pages=1590–1602 |doi=10.1890/03-5109 |jstor=4493673|bibcode=2004EcoAp..14.1590V }}</ref>
thumb|upright|Sugar pine starting to succumb to white pine blister rust
The species is generally resistant to fire because of its thick bark and because it clears away competing species.<ref name="Arno" /> However, its mortality has been directly linked to drier conditions and higher temperatures. Climate change presents a threat to species health: higher temperatures can decrease resin levels within the trees, weakening defenses against pathogens. At the same time, warmer winters increase survival of pests and pathogens. The weakened or dying trees then provide fuel for forest fires, which may become more frequent and more intense with rising summer temperatures, particularly if coupled with drier conditions and stronger winds.<ref>{{cite journal |last1=Slack |first1=A |last2=Kane |first2=J |last3=Knapp |first3=E |last4=Sherriff |first4=R |year=2017 |title=Contrasting impacts of climate and competition on large sugar pine growth and defense in a fire-excluded forest of the central sierra nevada |journal=Forests |volume=8 |issue=7 |page=244 |doi=10.3390/f8070244 |doi-access=free|bibcode=2017Fore....8..244S }}</ref>
=== Protective efforts === Sugar pine trees are in slow decline due to several threats; white pine blister rust, mountain pine beetles, and climate change. Efforts to restore sugar pines and other white pine trees that have been impacted by invasive species, climate change, and fires have been undertaken by governmental and non-governmental entities. One nonprofit, the Sugar Pine Foundation, was created in 2004 to plant sugar pine seeds in the Sierra Nevada along the border of California and Nevada.<ref name="spf" /> They plant seedlings grown from seeds collected from tree strains resistant to blister rust. The foundation's aim is to build in the a wild a sugar pine population that is resistant to white pine rust.<ref>{{cite journal |last1=Maloney |first1=PE |last2=Vogler |first2=DR |last3=Eckert |first3=AJ |last4=Jensen |first4=CE |last5=Neale |first5=DB |year=2011 |title=Population biology of sugar pine (''Pinus lambertiana'' Dougl.) with reference to historical disturbances in the Lake Tahoe Basin: Implications for restoration |journal=Forest Ecology and Management |volume=262 |issue=5 |pages=770–779 |doi=10.1016/j.foreco.2011.05.011|bibcode=2011ForEM.262..770M }}</ref>
==Uses== [[File:Sugar Pine Bark.jpg|thumb|right|upright|Bark of a sugar pine on Mount San Antonio in the San Gabriel Mountains of Southern California]]
According to David Douglas, who was guided to the (exceptionally thick) tree specimen he was looking for by a Native American,<ref name="Arno" /> some tribes ate the sweetish seeds. These were eaten raw and roasted, and also used to make flour or pulverized into a spread.<ref name="Arno" /> Native Americans also ate the inner bark.<ref name="Arno" /> The sweet sap or pitch was consumed, in small quantities due to its laxative properties,<ref name=Peattie>{{cite book |last=Peattie |first=Donald Culross |author-link=Donald C. Peattie |title=A Natural History of Western Trees |year=1953 |publisher=Bonanza Books |location=New York |page=55}}</ref> but could also be chewed as gum.<ref name="Arno" /> Its flavor is thought largely to be derived from the pinitol it contains.<ref name="Arno" />
In the mid-19th century, the trees were used liberally as lumber during the California Gold Rush. In modern times they are used in much lower quantities, being spared for high-end products as with Western white pine.<ref name="Arno" />
The odorless wood is also preferred for packing fruit, as well as storing drugs and other goods. Its straight grain also makes it a useful organ pipe material.<ref name=Peattie/> The wood was also long used for piano keys; in 1907 or 1908 the Connection piano-action maker Pratt, Read & Co. purchased "950,000 feet of clear sugar pine" for that use in & around Placerville, CA.<ref>"Buying Lumber for Keys". Music Trade Review 47:10 (5 September 1908), 35.</ref>
==Folklore== In the Achomawi creation myth, Annikadel, the creator, makes one of the 'First People' by intentionally dropping a sugar pine seed in a place where it can grow. One of the descendants in this ancestry is Sugarpine-Cone man, who has a handsome son named Ahsoballache.<ref name="Woiche">{{cite book |title=Annikadel: The History of the Universe as Told by the Achumawi Indians of California |last=Woiche |first=Istet |editor1-last=Merriam |editor1-first=Clinton Hart |editor1-link=Clinton Hart Merriam |year=1992 |publisher=University of Arizona Press |location=Tucson |isbn=978-0-8165-1283-6 |oclc=631716557 }}</ref>
After Ahsoballache marries the daughter of To'kis the Chipmunk-woman, his grandfather insists that the new couple have a child. To this end, the grandfather breaks open a scale from a sugar pine cone, and secretly instructs Ahsoballache to immerse the scale's contents in spring water, then hide them inside a covered basket. Ahsoballache performs the tasks that night; at the next dawn, he and his wife discover the infant Edechewe near their bed.<ref name="Woiche" />
The Washo language has a word for sugar pine, {{lang|was|simt'á:gɨm}}, and also a word for "sugar pine sugar", {{lang|was|nanómba}}.
==References== {{Reflist}}
==Further reading== * {{cite book | last1 = Chase| first1 = J. Smeaton| author-link1 = J. Smeaton Chase|others=Eytel, Carl (illustrations)|title = Cone-bearing Trees of the California Mountains | url = https://archive.org/details/conebearingtrees00chas/page/12/mode/2up| location = Chicago | publisher = A.C. McClurg & Co. | pages = 12–14 |year=1911 | oclc = 3477527|lccn=11004975}} * {{Silvics |first1=Bohun B. |last1=Kinloch Jr. |first2=William H. |last2=Scheuner |volume=1 |genus=Pinus |species=lambertiana}} * {{FEIS |last=Habeck |first=R. J. |date=1992 |type=tree |genus=Pinus |species=lambertiana }}
==External links== {{Commons}} * [http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?Pinus%20lambertiana U.C. Jepson Manual treatment for ''Pinus lambertiana''] * [http://www.fs.fed.us/r6/dorena/rust/index.shtml US Forest Service – Dorena Genetic Resource Center] – USFS rust resistance program * [http://www.sugarpinefoundation.org The Sugar Pine Foundation] – ''The Sugar Pine and Western White Pine Restoration Program'' * {{CalPhotos|Pinus|lambertiana}} * {{cite iucn |author=Farjon, A. |date=2013 |title=''Pinus lambertiana'' |volume=2013 |article-number=e.T42374A2976106 |doi=10.2305/IUCN.UK.2013-1.RLTS.T42374A2976106.en |access-date=13 November 2021}} * [http://www.pinetum.org/cones/PNlambertiana.jpg Arboretum de Villardebelle: photo of a cone]
{{Taxonbar|from=Q227779}} {{Authority control}}
lambertiana Category:Trees of Northern America Category:Flora of California Category:Flora of the Sierra Nevada (United States) Category:Natural history of the California Coast Ranges Category:Natural history of the Transverse Ranges Category:Edible nuts and seeds Category:Plants used in traditional Native American medicine Category:Least concern flora of the United States