View source for Biology:Spruce

{{Short description|Genus of evergreen, coniferous tree}}
{{Automatic taxobox
| name = Spruce
| fossil_range = {{fossil range|Valanginian|Recent}}
| image = Picea abies.jpg
| image_caption = Norway spruce (''Picea abies'')
| parent_authority = Frankis
| taxon = Picea
| type_species = ''[[Biology:Picea abies|Picea abies]]''
| type_species_authority = ([[Biography:Carl Linnaeus|L.]]) [[Biography:Gustav Karl Wilhelm Hermann Karsten|H. Karst.]]
| authority = Mill.
| subdivision_ranks = Species
| subdivision = About 35; see text.
| synonyms = 
* ''Veitchia'' <small>Lindley</small>
}}

A '''spruce''' is a [[Earth:Tree|tree]] of the genus '''''Picea''''' ({{IPAc-en|p|aɪ|ˈ|s|iː|.|ə}} {{respell|py|SEE|ə}}),<ref>''Sunset Western Garden Book,'' 1995:606–607</ref> a genus of about 35 species of [[Biology:Conifer|conifer]]ous [[Biology:Evergreen|evergreen]] trees in the family [[Biology:Pinaceae|Pinaceae]], found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfamily '''Piceoideae'''. Spruces are large trees, from about 20 to 60 m (about 60&ndash;200&nbsp;ft) tall when mature, and have [[Biology:Whorl (botany)|whorled]] branches and conical form. 

They can be distinguished from other members of the pine family by their needles (leaves), which are four-sided and attached singly to small persistent peg-like structures (pulvini or sterigmata<ref>Hart, G.T. (2018) ''Plants in Literature and Life: a wide-ranging dictionary of botanical terms''. FriesenPress. {{ISBN|978-1770674417}}</ref>) on the branches, and by their cones (without any protruding [[Biology:Bract|bract]]s), which hang downwards after they are pollinated.<ref>{{cite web |title=Pine Relatives |url=http://www.lovettpinetum.org/pines/pine-relatives/ |publisher=Lovett Pinetum |access-date=2018-08-27}}</ref> The needles are shed when 4–10 years old, leaving the branches rough with the retained pegs. In other similar genera, the branches are fairly smooth.

Spruce are used as food plants by the [[Biology:Larva|larva]]e of some [[Biology:Lepidoptera|Lepidoptera]] (moth and butterfly) species, such as the [[Biology:Choristoneura fumiferana|eastern spruce budworm]]. They are also used by the larvae of [[Biology:Gall adelgid|gall adelgid]]s (''Adelges'' species).

In the mountains of western Sweden, scientists have found a Norway spruce, nicknamed Old Tjikko, which by reproducing through [[Biology:Layering|layering]], has reached an age of 9,550 years and is claimed to be the world's oldest known living tree.<ref>{{cite web |url=http://www.sciam.com/podcast/episode.cfm?id=2AA00080-0AD4-55E1-EA50BC7A36A218C4& |title=Swedish Spruce Is World's Oldest Tree: Scientific American Podcast |date=27 May 2008 |access-date=21 August 2016}}</ref>

== Description ==

===Morphology===
[[File:Picea abies Nadelkissen.jpg|thumb|The peg-like base of the needles, or pulvinus, in Norway spruce (''Picea abies'').]]
[[File:Picea glauca twig Denali NP AK.jpg|thumb|Pulvini remain after the needles fall (white spruce, ''Picea glauca'').]]
Determining that a tree is a spruce is not difficult; [[Biology:Evergreen|evergreen]] needles that are more or less quadrangled, and especially the [[Biology:Pulvinus|pulvinus]], give it away. Beyond that, determination can become more difficult. Intensive sampling in the Smithers/Hazelton/Houston area of British Columbia showed Douglas (1975),<ref name="doug">Douglas, G.W. (1975). Spruce (''Picea'') hybridization in west-central British Columbia. B.C. Min. For., Forest Science, Smithers BC, unpublished report, cited by Coates et al. 1994. (Cited by Coates et al. 1994, orig. not seen)</ref> according to Coates et al. (1994),<ref name="coat">Coates, K.D.; Haeussler, S.; Lindeburgh, S.; Pojar, R.; Stock, A.J. (1994). Ecology and silviculture of interior spruce in British Columbia. Canada/British Columbia Partnership Agreement For. Resour. Devel., Victoria BC, FRDA Rep. 220. 182 p.</ref> that cone scale [[Biology:Morphology|morphology]] was the feature most useful in differentiating species of spruce; the length, width, length: width ratio, the length of free scale (the distance from the imprint of the seed wing to the tip of the scale), and the percentage free scale (length of free scale as a percentage of the total length of the scale) were most useful in this regard. Daubenmire (1974),<ref name="daub">{{cite journal|author=Daubenmire, R. |year=1974|title=Taxonomic and ecologic relationships between ''Picea glauca'' and ''Picea engelmannii''|journal= Can. J. Bot. |volume=52|issue=7|pages=1545–1560|doi=10.1139/b74-203}}</ref> after range-wide sampling, had already recognized the importance of the 2 latter characters. Taylor (1959)<ref name="taylor">{{cite journal|author=Taylor, T.M.C. |year=1959|title= The taxonomic relationship between ''Picea glauca'' (Moench) Voss and ''P. engelmannii'' Parry|journal= Madroño |jstor=41422994|volume=15|issue=4|pages=111–115}} (Cited in Coates et al. 1994).</ref> had noted that the most obvious morphological difference between typical ''Picea glauca'' and typical ''P. engelmannii'' was the cone scale, and Horton (1956,1959)<ref name="horton1">Horton, K.W. (1956). A taxonomic and ecological study of ''Picea glauca'' and ''Picea engelmannii'' in North America. Diploma thesis, Oxford Univ., U.K. 103 p.</ref><ref name="horton2">Horton, K.W. (1959). Characteristics of subalpine spruce in Alberta. Can. Dep. Northern Affairs National Resour., For. Branch, For. Res. Div., Ottawa ON, Tech. Note 76. 20 p.</ref> found that the most useful diagnostic features of the 2 spruces are in the cone; differences occur in the flower, shoot and needle, "but those in the cone are most easily assessed" (Horton 1959).<ref name="horton2" /> Coupé et al. (1982)<ref name="coup">Coupé, R.; Ray, C.A.; Comeau, A.; Ketcheson, M.V.; Annas, R.M. (1982). A guide to some common plants of the Skeena area, British Columbia. B.C. Min. For., Res. Branch, Victoria BC.</ref> recommended that cone scale characters be based on samples taken from the midsection of each of 10 cones from each of 5 trees in the population of interest.

Without cones, morphological differentiation among spruce species and their hybrids is more difficult. Species classification for seeds collected from spruce stands in which introgressive hybridization between white and Sitka spruces (''P. sitchensis'') may have occurred is important for determining appropriate cultural regimens in the nursery. If, for instance, white spruce grown at container nurseries in southwestern British Columbia are not given an extended photoperiod, leader growth ceases early in the first growing season, and seedlings do not reach the minimum height specifications.<ref name="arn1">Arnott, J.T. (1974). "Germination and seedling establishment". pp. 55–66 in Cayford, J.H. (Ed.). ''Direct Seeding Symposium'', Timmins ON, Sept. 1973, Can. Dep. Environ., Can. For. Serv., Ottawa ON, Proc., Publ. 1339.</ref><ref name="arn2">{{cite journal | last1 = Arnott | first1 = J.T. | year = 1979 | title = Effect of light intensity during extended photoperiod on growth of amabalis fir, mountain hemlock, and white and Engelmann spruce seedlings | journal = Can. J. For. Res. | volume = 9 | pages = 82–89 | doi=10.1139/x79-014}}</ref> But, if an extended photoperiod is provided for Sitka spruce, seedlings become unacceptably tall by the end of the first growing season <!-- (Arnott, unpublished data, cited by Yeh and Arnott 1986) -->.<ref name="yeh" /> Species classification of seedlots collected in areas where hybridization of white and Sitka spruces has been reported has depended on (i) easily measured cone scale characters of seed trees, especially free scale length, (ii) visual judgements of morphological characters, e.g., growth rhythm, shoot and root weight, and needle serration, or (iii) some combination of (i) and (ii) (Yeh and Arnott 1986).<ref name="yeh">{{cite journal | last1 = Yeh | first1 = F.C. | last2 = Arnott | first2 = J.T. | year = 1986 | title = Electrophoretic and morphological differentiation of ''Picea sitchensis, Picea glauca,'' and their hybrids | journal = Can. J. For. Res. | volume = 16 | issue = 4| pages = 791–798 | doi = 10.1139/x86-140}}</ref> Useful to a degree, these classification procedures have important limitations; genetic composition of the seeds produced by a stand is determined by both the seed trees and the pollen parents, and species classification of hybrid seedlots and estimates of their level of introgression on the basis of seed-tree characteristics can be unreliable when hybrid seedlots vary in their introgressiveness in consequence of spatial and temporal variations in contributions from the pollen parent (Yeh and Arnott 1986).<ref name="yeh" /> Secondly, morphological characters are markedly influenced by ontogenetic and environmental influences, so that to discern spruce hybrid seedlot composition with accuracy, hybrid seedlots must differ substantially in morphology from both parent species. Yeh and Arnott (1986)<ref name="yeh" /> pointed out the difficulties of estimating accurately the degree of introgression between white and Sitka spruces; introgression may have occurred at low levels, and/or hybrid seed lots may vary in their degree of introgression in consequence of repeated backcrossing with parental species.

=== Growth ===
Spruce seedlings are most susceptible immediately following [[Biology:Germination|germination]], and remain highly susceptible through to the following spring. More than half of spruce seedling mortality probably occurs during the first growing season and is also very high during the first winter,<ref name="alex">Alexander, R.R. (1987). Ecology, silviculture, and management of the Engelmann spruce–subalpine fir type in the central and southern Rocky Mountains. USDA, For. Serv., Washington DC, Agric. Handb. 659. 144 p.</ref> when seedlings are subjected to freezing damage, frost heaving and erosion, as well as smothering by litter and snow-pressed vegetation. Seedlings that germinate late in the growing season are particularly vulnerable because they are tiny and have not had time to [[Earth:Hardiness (plants)|harden off]] fully.

Mortality rates generally decrease sharply thereafter, but losses often remain high for some years. "Establishment" is a subjective concept based on the idea that once a seedling has successfully reached a certain size, not much is likely to prevent its further development. Criteria vary, of course, but Noble and Ronco (1978),<ref name="nob">Noble, D.L.; Ronco, F. (1978). Seedfall and establishment of Engelmann spruce and subalpine fir in clearcut openings in Colorado. USDA, For. Serv., Rocky Mountain For. Range Exp. Sta., Res. Pap. RM-200. 12 p.</ref> for instance, considered that seedlings four to five years old, or 8&nbsp;cm to 10&nbsp;cm tall, warranted the designation "established", since only unusual factors such as [[Biology:Phacidiaceae|snow mold]], [[Biology:Wildfire|fire]], [[Earth:Disturbance (ecology)|trampling]], or predation would then impair regeneration success. Eis (1967)<ref name="eis">{{cite journal|author=Eis, S. |year=1967|title= Establishment and early development of white spruce in the interior of British Columbia|journal= For. Chron.|volume= 43|issue=2|pages=174–177|doi=10.5558/tfc43174-2|doi-access=free}}</ref> suggested that in dry habitats on either mineral soil or litter seedbeds a 3-year-old seedling may be considered established; in moist habitats, seedlings may need 4 or 5 years to become established on mineral soil, possibly longer on litter seedbeds.

Growth remains very slow for several to many years. Three years after shelterwood felling in subalpine Alberta, dominant [[Biology:Regeneration|regeneration]] averaged 5.5&nbsp;cm in height in scarified blocks, and 7.3&nbsp;cm in non-scarified blocks (Day 1970),<ref name="day2">Day, M.W.; Rudolph, V.J. (1970). Development of a white spruce plantation. Michigan State Univ., Agric. Exp. Sta., East Lansing MI, Res. Pap. 111. 4 p.</ref> possibly reflecting diminished fertility with the removal of the A horizon.

== Taxonomy ==

=== Classification ===
[[Biology:DNA|DNA]] analyses have shown that traditional classifications based on the [[Biology:Morphology|morphology]] of needle and [[Conifer cone|cone]] are artificial.<ref name="Ran">{{cite journal |author1=Jin-Hua Ran |author2=Xiao-Xin Wei |author3=Xiao-Quan Wang |year=2006 |title=Molecular phylogeny and biogeography of ''Picea'' (Pinaceae): implications for phylogeographical studies using cytoplasmic haplotypes |url=http://lseb.ibcas.ac.cn/oldzjxx/wangxq/pdf/Picea.pdf |url-status=dead |journal=[[Biology:Molecular Phylogenetics and Evolution|Molecular Phylogenetics and Evolution]] |volume=41 |issue=2 |pages=405–419 |doi=10.1016/j.ympev.2006.05.039 |pmid=16839785 |archive-url=https://web.archive.org/web/20120424225243/http://lseb.ibcas.ac.cn/oldzjxx/wangxq/pdf/Picea.pdf |archive-date=24 April 2012}}</ref><ref name="Sigurgeirsson">{{cite journal |author1=Aðalsteinn Sigurgeirsson |author2=Alfred E. Szmidt |name-list-style=amp |year=1993 |title=Phylogenetic and biogeographic implications of chloroplast DNA variation in ''Picea'' |journal=Nordic Journal of Botany |volume=13 |issue=3 |pages=233–246 |doi=10.1111/j.1756-1051.1993.tb00043.x}}</ref> A 2006 study found that ''[[Biology:Picea breweriana|P. breweriana]]'' had a [[Biology:Basal (phylogenetics)|basal]] position, followed by ''[[Biology:Picea sitchensis|P. sitchensis]]'',<ref name="Ran" /> and the other species were further divided into three [[Biology:Clade|clade]]s, suggesting that ''Picea'' originated in North America. The oldest record of spruce that has been found in the fossil record is from the Early Cretaceous ([[Earth:Valanginian|Valanginian]]) of western Canada, around 136 million years old.<ref>{{cite journal |author1=Ashley A. Klymiuk |author2=Ruth A. Stockey |name-list-style=amp |year=2012 |title=A Lower Cretaceous (Valanginian) seed cone provides the earliest fossil record for Picea (Pinaceae) |journal=American Journal of Botany |volume=99 |issue=6 |pages=1069–1082 |doi=10.3732/ajb.1100568 |pmid=22623610 |doi-access=free}}</ref>

===Species===
{{cladogram|title=Phylogeny of ''Picea''<ref>{{cite journal |last1=Stull |first1=Gregory W. |last2=Qu |first2=Xiao-Jian |last3=Parins-Fukuchi |first3=Caroline |last4=Yang |first4=Ying-Ying |last5=Yang |first5=Jun-Bo |last6=Yang |first6=Zhi-Yun |last7=Hu |first7=Yi  |last8=Ma |first8=Hong |last9=Soltis |first9=Pamela S. |last10=Soltis |first10=Douglas E. |last11=Li |first11=De-Zhu |last12=Smith |first12=Stephen A. |last13=Yi |first13=Ting-Shuang |display-authors=et al. |year=2021 |title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms |journal=Nature Plants |url=https://www.nature.com/articles/s41477-021-00964-4 |volume=7 |issue= 8|pages=1015–1025 |doi=10.1038/s41477-021-00964-4|biorxiv=10.1101/2021.03.13.435279 |pmid= 34282286|pmc= |bibcode= |s2cid=232282918 |doi-access=}}</ref><ref>{{cite journal |last1=Stull |first1=Gregory W. |display-authors=et al. |year=2021 |title=main.dated.supermatrix.tree.T9.tre |publisher=Figshare |doi=10.6084/m9.figshare.14547354.v1 |url=https://figshare.com/articles/dataset/Gene_duplications_and_genomic_conflict_underlie_major_pulses_of_phenotypic_evolution_in_gymnosperms/14547354 |doi-access=}}</ref>|
{{clade|style=font-size:90%;line-height:100%;width:400px
|1={{clade
  |1={{clade
    |label1=section
    |sublabel1=''Sitcha''
    |1=''[[Biology:Picea sitchensis|P. sitchensis]]'' <small>(Bongard) Carrière</small>
     }}
  |2={{clade
    |1={{clade
      |label1=section
      |sublabel1=''Brewerianae''
      |1=''[[Biology:Picea breweriana|P. breweriana]]'' <small>Watson</small>
       }}
    |2={{clade
      |label1=section
      |sublabel1=''Casicta''
      |1={{clade
        |1={{clade
          |label1=series
          |sublabel1=''Likiangenses''
          |1={{clade
            |1=''[[Biology:Picea likiangensis|P. likiangensis]]'' <small>(Franchet) Pritzel</small>
            |2={{clade
              |1=''[[Biology:Picea farreri|P. farreri]]'' <small>Page & Rushforth</small>
              |2=''[[Biology:Picea spinulosa|P. spinulosa]]'' <small>(Griffith) Henry</small>
               }}
             }}
           }}
        |2={{clade
          |1={{clade
            |label1=series
            |sublabel1=''Schrenkianae''
            |1=''[[Biology:Picea schrenkiana|P. schrenkiana]]'' <small>Fischer & Meyer</small>
             }}
          |2={{clade
            |label1=series
            |sublabel1=''Glaucae''
            |1={{clade
              |1=''[[Biology:Picea smithiana|P. smithiana]]'' <small>(Wallich) Boiss.</small>
              |2={{clade
                |1=''[[Biology:Picea glauca|P. glauca]]'' <small>(Moench) Voss</small>
                |2=''[[Biology:Picea engelmannii|P. engelmannii]]'' <small>Parry ex Engelmann</small>
                 }}
               }}
            |label2=series
            |sublabel2=''Orientales''
            |2={{clade
              |1={{clade
                |1=''[[Biology:Picea martinezii|P. martinezii]]'' <small>T.F.Patt.</small>
                |2={{clade
                  |1=''[[Biology:Picea chihuahuana|P. chihuahuana]]'' <small>Martínez</small>
                  |2=''[[Biology:Picea alcoquiana|P. alcoquiana]]'' <small>(Veitch ex Lindley) Carrière</small>
                   }}
                 }}
              |2={{clade
                |1={{clade
                  |1=''[[Biology:Picea brachytyla|P. brachytyla]]'' <small>(Franchet) Pritzel</small>
                  |2={{clade
                    |1=''[[Biology:Picea neoveitchii|P. neoveitchii]]'' <small>Masters</small>
                    |2=''[[Biology:Picea morrisonicola|P. morrisonicola]]'' <small>Hayata</small>
                     }}
                   }}
                |2={{clade
                  |1={{clade
                    |1=''[[Biology:Picea purpurea|P. purpurea]]'' <small>Masters</small>
                    |2=''[[Biology:Picea wilsonii|P. wilsonii]]'' <small>Masters</small>
                     }}
                  |2={{clade
                    |1=''[[Biology:Picea orientalis|P. orientalis]]'' <small>(von Linné) Peterm.</small>
                    |2={{clade
                      |1=''[[Biology:Picea maximowiczii|P. maximowiczii]]'' <small>Regel ex Masters</small>
                      |2=''[[Biology:Picea polita|P. polita]]'' <small>(Siebold & Zuccarini) Carrière</small>
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
      |label2=section
      |sublabel2=''Picea''
      |2={{clade
        |label1=series
        |sublabel1=''Pungentes''
        |1=''[[Biology:Picea pungens|P. pungens]]'' <small>Engelmann</small>
        |2={{clade
          |label1=series
          |sublabel1=''Jezoenses''
          |1={{clade
            |1={{clade
              |1=''[[Biology:Picea glehnii|P. glehnii]]'' <small>(Schmidt) Masters</small>
              |2=''[[Biology:Picea jezoensis|P. jezoensis]]'' <small>(Sieb. & Zuccarini) Carrière</small>
               }}
            |2={{clade
              |1=''[[Biology:Picea rubens|P. rubens]]'' <small>Sargent</small>
              |2={{clade
                |1=''[[Biology:Picea mariana|P. mariana]]'' <small>(Miller) Britton, Sterns & Poggenburg</small>
                |2=''[[Biology:Picea omorika|P. omorika]]'' <small>(Panèiæ) Purkyne</small>
                 }}
               }}
             }}
          |label2=series
          |sublabel2=''Abies''
          |2={{clade
            |1=''[[Biology:Picea obovata|P. obovata]]'' <small>Ledeb.</small>
            |2={{clade
              |1={{clade
                |1=''[[Biology:Picea abies|P. abies]]'' <small>(von Linné) Karsten</small>
                |2=''[[Biology:Picea koyamae|P. koyamae]]'' <small>Shiras.</small>
                 }}
              |2={{clade
                |1=''[[Biology:Picea asperata|P. asperata]]'' <small>Masters</small>
                |2={{clade
                  |1=''[[Biology:Picea koraiensis|P. koraiensis]]'' <small>Nakai</small>
                  |2={{clade
                    |1=''[[Biology:Picea torano|P. torano]]'' <small>(Siebold ex Koch) Koehne</small>
                    |2={{clade
                      |1=''[[Biology:Picea retroflexa|P. retroflexa]]'' <small>Masters</small>
                        |2={{clade
                          |1=''P. shirasawae'' <small>Hayashi</small>
                          |2={{clade
                            |1=''[[Biology:Picea crassifolia|P. crassifolia]]'' <small>Komarov</small>
                            |2=''[[Biology:Picea meyeri|P. meyeri]]'' <small>Rehder & Wilson</small>
                             }}
                           }}
                         }}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}

{{As of|2022|April}}, [[Biology:Plants of the World Online|Plants of the World Online]] accepted 37 species.<ref name="POWO_30005648-2">{{cite web |title=''Picea'' A.Dietr. |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30005648-2 |access-date=2022-04-20 }}</ref> The grouping is based on Ran et al. (2006).<ref name="Ran"/>
*Basal species:
** ''[[Biology:Picea breweriana|Picea breweriana]]'' – Brewer's spruce, Klamath Mountains, North America; local endemic
** ''[[Biology:Picea sitchensis|Picea sitchensis]]'' – Sitka spruce, Pacific coast of North America; the largest species, to 95&nbsp;m tall; important in forestry
*Clade I (Northern and western [[Earth:North America|North America]], in boreal forests or high mountains)
** ''[[Biology:Picea engelmannii|Picea engelmannii]]'' – Engelmann spruce, western North American mountains; important in forestry
** ''[[Biology:Picea glauca|Picea glauca]]'', syn. ''Picea laxa'' – white spruce, northern North America; important in forestry
*Clade II (throughout Asia, mostly in mountainous areas, a few isolated populations in higher elevations of Mexico,)
** ''[[Biology:Picea brachytyla|Picea brachytyla]]'' – Sargent's spruce, southwest {{wipe|China}}
** ''[[Biology:Picea chihuahuana|Picea chihuahuana]]'' – Chihuahua spruce, northwest Mexico (rare)
** ''[[Biology:Picea farreri|Picea farreri]]'' – Burmese spruce, northeast Burma, southwest {{wipe|China}} (mountains)
** ''[[Biology:Picea likiangensis|Picea likiangensis]]'' – Likiang spruce, southwest China
** ''[[Biology:Picea martinezii|Picea martinezii]]'' – Martinez spruce, northeast Mexico (very rare, endangered)
** ''[[Biology:Picea maximowiczii|Picea maximowiczii]]'' – Maximowicz spruce, {{wipe|Japan}} (rare, mountains)
** ''[[Biology:Picea morrisonicola|Picea morrisonicola]]'' – Taiwan spruce, Taiwan (high mountains)
** ''[[Biology:Picea neoveitchii|Picea neoveitchii]]'' – Veitch's spruce, northwest China (rare, endangered)
** ''[[Biology:Picea orientalis|Picea orientalis]]'' – Caucasian spruce or Oriental spruce, [[Earth:Caucasus|Caucasus]], northeast [[Place:Turkey|Turkey]]
** ''[[Biology:Picea polita|Picea polita]]'', syn. ''Picea torano'' – tiger-tail spruce, Japan
** ''[[Biology:Picea purpurea|Picea purpurea]]'' – purple cone spruce, western China
** ''[[Biology:Picea schrenkiana|Picea schrenkiana]]'' – Schrenk's spruce, mountains of central Asia
** ''[[Biology:Picea smithiana|Picea smithiana]]'' – morinda spruce, western {{wipe|Himalaya}}, eastern Afghanistan, northern and northwest {{wipe|India}}
** ''[[Biology:Picea spinulosa|Picea spinulosa]]'' – Sikkim spruce, northeast India (Sikkim), eastern {{wipe|Himalaya}}
** ''[[Biology:Picea wilsonii|Picea wilsonii]]'' – Wilson's spruce, western China
*Clade III (Europe, Asia, and North America, mostly in boreal forests or mountainous areas)
** ''[[Biology:Picea abies|Picea abies]]'' – Norway spruce, Europe; important in forestry, the original Christmas tree
** ''[[Biology:Picea alcoquiana|Picea alcoquiana]]'' – ("''P. bicolor''") Alcock's spruce, central Japan (mountains)
** ''[[Biology:Picea asperata|Picea asperata]]'' – dragon spruce, western China; several varieties
** ''[[Biology:Picea crassifolia|Picea crassifolia]]'' – Qinghai spruce, China
** ''[[Biology:Picea glehnii|Picea glehnii]]'' – Glehn's spruce, northern Japan, Sakhalin
** ''[[Biology:Picea jezoensis|Picea jezoensis]]'' – Jezo spruce, northeast Asia, Kamchatka south to Japan
** ''[[Biology:Picea koraiensis|Picea koraiensis]]'' – Korean spruce, [[Place:Korea|Korea]], northeast China
** ''[[Biology:Picea koyamae|Picea koyamae]]'' – Koyama's spruce, Japan (mountains)
** ''[[Biology:Picea mariana|Picea mariana]]'' – black spruce, northern North America
** ''[[Biology:Picea meyeri|Picea meyeri]]'' – Meyer's spruce, northern China (from Inner Mongolia to Gansu)
** ''[[Biology:Picea obovata|Picea obovata]]'' – Siberian spruce, north [[Place:Scandinavia|Scandinavia]], Siberia; often treated as a variant of ''P. abies'' (and hybridises with it), but has distinct cones
** ''[[Biology:Picea omorika|Picea omorika]]'' – Serbian spruce, Serbia and Bosnia; local endemic; important in horticulture
** ''[[Biology:Picea pungens|Picea pungens]]'' – blue spruce or Colorado spruce, Rocky Mountains, North America; important in horticulture
** ''[[Biology:Picea retroflexa|Picea retroflexa]]'' – green dragon spruce, China
** ''[[Biology:Picea rubens|Picea rubens]]'' – red spruce, northeastern North America; important in forestry, known as [[Biology:Picea rubens|Adirondack]] in musical-instrument making
*Others
** ''[[Biology:Picea aurantiaca|Picea aurantiaca]]'' <small>Mast.</small>
** ''Picea austropanlanica'' <small>Silba</small>
** ''Picea linzhiensis'' <small>(W.C.Cheng & L.K.Fu) Rushforth</small>
*Hybrids
** ''Picea'' × ''albertiana'' <small>S.Br.</small>
** ''Picea'' × ''fennica'' <small>(Regel) Kom.</small>
** [[Biology:Picea × lutzii|''Picea'' × ''lutzii'']] <small>Little</small>
** ''Picea'' × notha'' <small>Rehder</small>

==Fossil species==
There are also a number of extinct species identified from fossil evidence:
===Cones===
{{div col}}
*†''Picea anadyrensis'' {{small|Kryshtofovich}}
*†''Picea antiqua'' {{small|Vasskovsky}}
*†''Picea beckii'' {{small|Mai}}
*†''Picea bilibinii'' {{small|Vasskovsky}}
*†''Picea burtonii'' {{small|Klymiuk & Stockey}}
*†''Picea camtschatica'' {{small|Vasskovsky}}
*†''Picea deweyensis'' {{small|Axelrod}}
*†''Picea diettertiana'' {{small|Miller}}
*†''Picea eichhornii'' {{small|Miller}}
*†''Picea evenica'' {{small|Vasskovsky}}
*†''Picea fimbriata'' {{small|Chelebaeva}}
*†''Picea garoensis'' {{small|Tanai & Suzuki}}
*†''Picea harrimani'' {{small|Knowlton}}
*†''Picea hondoensis'' {{small|Vasskovsky}}
*†''Picea indigirensis'' {{small|Peresvetov}}
*†''Picea latibracteata'' {{small|Miki}}
*†''Picea latisquamosa'' {{small|Kinkelin}}
*†''Picea metechensis'' {{small|Kharatishvili}}
*†''Picea mioorientalis'' {{small|Usnadze}}
*†''Picea mugodzharica'' {{small|Rajushkina}}
*†''Picea oligocaenica'' {{small|Engelhardt}}
*†''Picea pevekensis'' {{small|Golovneva}}
*†''Picea praeajanensis'' {{small|Vasskovsky}}
*†''Picea protopicea'' {{small|(Velenovský) Tyroff}}
*†''Picea sookensis'' {{small|LaMotte}}
*†''Picea sugaii'' {{small|Tanai & Onoe}}
*†''Picea suifunensis'' {{small|Kryshtofovich}}
*†''Picea vassiljevii'' {{small|Vasskovsky}}
*†''Picea vitjasii'' {{small|Vasskovsky}}
*†''Picea wolfei'' {{small|Crabtree}}
*†''Picea wollosowiczii'' {{small|Sukaczev}}
*†''Picea yanensis'' {{small|Golovneva & Shczepetov}}
{{div col end}}

===Foliage===
{{div col}}
*†''Picea cretacea'' {{small|Velenovský}}
*†''Picea echinata'' {{small|Müller-Stoll}}
*†''Picea korfiensis'' {{small|Chelebaeva}}
*†''Picea lahontensis'' {{small|MacGinitie}}
*†''Picea morosovae'' {{small|Akhmetiev & Shevyreva}}
*†''Picea nakauchii'' {{small|Matsumoto, Ohsawa, & Nishida}}
*†''Picea palaeomorika'' {{small|Müller-Stoll}}
*†''Picea quilchensis'' {{small|Penhallow}}
*†''Picea tranquillensis'' {{small|Penhallow}}
{{div col end}}

===Pollen===
{{div col}}
*†''Picea alata'' {{small|Zaklinsk}}
*†''Picea bella'' {{small|Bolkhovitina}}
*†''Picea complanatiformis'' {{small|Bolkhovitina}}
*†''Picea depressa'' {{small|Bolkhovitina}}
*†''Picea distorta'' {{small|Bolkhovitina}}
*†''Picea distracta'' {{small|Bolkhovitina}}
*†''Picea exilioides'' {{small|Bolkhovitina}}
*†''Picea gigantissima'' {{small|Bolkhovitina}}
*†''Picea grandipollinia'' {{small|Ananova}}
*†''Picea grandis'' {{small|Panova}}
*†''Picea grandivescipites'' {{small|Wodehouse}}
*†''Picea kryshtofovichii'' {{small|Ammosov}}
*†''Picea longisaccata'' {{small|Rovnina}}
*†''Picea media'' {{small|Ananova}}
*†''Picea mesophytica'' {{small|Pokrovskaja}}
*†''Picea minor'' {{small|Mtchedlishvili}}
*†''Picea multigruma'' {{small|Chlonova}}
*†''Picea omoriciformis'' {{small|Bolkhovitina}}
*†''Picea parvireticulata'' {{small|Rovnina}}
*†''Picea pseudorotundiformis'' {{small|Maljavkina}}
*†''Picea rara'' {{small|Stanley}}
*†''Picea sacculifera'' {{small|(Maljavkina) Chlonova}}
*†''Picea samoilovitchiana'' {{small|Rovnina}}
*†''Picea schrenkianiformis'' {{small|Zaklinskaja}}
*†''Picea scotica'' {{small|Simpson}}
*†''Picea singularis'' {{small|Bolkhovitina}}
*†''Picea spirelliformis'' {{small|(Maljavkina) Bolkhovitina}}
*†''Picea sutschanensis'' {{small|Verbizkaja}}
*†''Picea tasaranica'' {{small|Zaklinskaja}}
*†''Picea tobolica'' {{small|Panova}}
*†''Picea valanjinica'' {{small|Rovnina}}
{{div col end}}

===Seeds===
{{div col}}
*†''Picea altaica'' {{small|Rajushkina}}
*†''Picea hiyamensis'' {{small|Tanai & Suzuki}}
*†''Picea kaneharae'' {{small|Tanai & Onoe}}
*†''Picea kanoi'' {{small|Huzioka}}
*†''Picea magna'' {{small|MacGinitie}}
*†''Picea pinifructus'' {{small|Brown}}
*†''Picea sonomensis'' {{small|Axelrod}}
{{div col end}}

===Wood===
*†''Picea palaeomaximowiczii'' {{small|Watari}}
*†''Picea wakimizui'' {{small|(Watari) Watari}}
*†''Picea withamii'' {{small|(Lindley & Hutton) Morris}}

===Multiple organs===
{{div col}}
*†''Picea columbiensis'' {{small|Penhallow}}
*†''[[Biology:Picea critchfieldii|Picea critchfieldii]]'' {{small|Jackson & Weng}} - Late Quaternary North America.<ref>{{Cite journal |last1=Jackson |first1=Stephen T. |last2=Weng |first2=Chengyu |date=1999-11-23 |title=Late Quaternary extinction of a tree species in eastern North America |journal=Proceedings of the National Academy of Sciences |language=en |volume=96 |issue=24 |pages=13847–13852 |bibcode=1999PNAS...9613847J |doi=10.1073/pnas.96.24.13847 |issn=0027-8424 |pmc=24153 |pmid=10570161 |doi-access=free}}</ref>
*†''Picea farjonii'' {{small|Herrera ''et al.''}}
*†''Picea heisseana'' {{small|von Fritsch}}
*†''Picea koribae'' {{small|Miki}}
*†''Picea miocenica'' {{small|Tanai}}
*†''Picea rotundosquamosa'' {{small|(Ludwig) Mai & Walther}}
*†''Picea ugoana'' {{small|Huzioka}}
{{div col end}}

===Unspecified===
*†''Picea alba'' {{small|Link}}
*†''Picea albertensis'' {{small|Penhallow}}
*†''Picea snatolensis'' {{small|Chelebaeva}}


===Fossil species formerly placed in ''Picea''===
*†''Picea cliffwoodensis'' {{small|Berry}} moved to †''Pityostrobus cliffwoodensis'' {{small|(Bery) Miller}}
*†''Picea succinifera'' {{small|(Göppert & Berendt)}} moved to ''Pinus succinifera'' {{small|(Göppert) Conwentz}}

== Etymology ==
[[File:Kuhmo.vaakuna.svg|thumb|upright=0.86|''Picea'' used in coat-of-arms of Kuhmo, Finland]]
''Spruce'', '' '' (1412), and ''{{not a typo|Sprws}} ve been generic terms for commodities brought to England by [[Organization:Hanseatic League|Hanseatic]] merchants (especially beer, boards, wooden chests and leather), and the tree thus was believed to be particular to Prussia, which for a time was figurative in England as a land of luxuries. It can be argued that the word is actually derived from the [[Social:Old French|Old French]] term {{lang|fro|Pruce}}, meaning literally Prussia.<ref>{{cite web |title=spruce - Origin and meaning of spruce by Online Etymology Dictionary |url=https://www.etymonline.com/word/spruce#etymonline_v_21904 |access-date=14 July 2020 |website=www.etymonline.com}}</ref>

== Ecology ==

=== Diseases ===
[[File:Fichtenstamm entrindet.jpg|thumb|Manually decorticated trunk of a spruce as protection against bark beetles]]
[[File:Древесина ели.jpg|thumb|Structure of spruce tree cells]]

==== ''Sirococcus'' blight (Deuteromycotina, Coelomtcetes) ====
The closely related species ''Sirococcus conigenus'' and ''Sirococcus piceicola'' cause shoot blight and seedling mortality of [[Biology:Pinophyta|conifers]] in North America, Europe, and North Africa.<ref>{{cite journal |last1=Rossman |first1=AY |last2=Castlebury |first2=LA |last3=Farr |first3=DF |last4=Stanosz |first4=GR |date=2008 |title=''Sirococcus conigenus'', ''Sirococcus piceicola'' sp. nov. and ''Sirococcus tsugae'' sp. nov. on conifers: anamorphic fungi in the Gnomoniaceae, Diaporthales |journal=Forest Pathology |volume=38 |pages=47–60 |doi=10.1111/j.1439-0329.2007.00529.x|doi-access=free }}</ref> Twig blight damage to seedlings of white and [[Biology:Picea rubens|red spruces]] in a nursery near Asheville, North Carolina, was reported by Graves (1914).<ref name="graves">{{cite journal |author=Graves, A.H. |year=1914 |title=Notes on diseases of trees in the southern Appalachians |journal=Phytopath |volume=4 |pages=63–72}}</ref> Hosts include white, [[Biology:Picea mariana|black]], [[Biology:Picea engelmannii|Engelmann]], [[Biology:Picea abies|Norway]], and [[Biology:Picea rubens|red spruces]], although they are not the plants most commonly damaged. ''Sirococcus'' blight of spruces in nurseries show up randomly in [[Biology:Seedling|seedling]]s to which the fungus was transmitted in infested seed. First-year seedlings are often killed, and larger plants may become too deformed for planting. Outbreaks involving < 30% of spruce seedlings in seedbeds have been traced to seed lots in which only 0.1% to 3% of seeds were infested. Seed infestation has in turn been traced to the colonization of spruce cones by ''S. conigenus'' in forests of the western interior. Infection develops readily if conidia are deposited on succulent plant parts that remain wet for at least 24 hours at 10&nbsp;°C to 25&nbsp;°C. Longer periods of wetness favour increasingly severe disease. Twig tips killed during growth the previous year show a characteristic crook.

==== ''Rhizosphaera kalkhoffi'' needle cast ====
''Rhizosphaera'' infects white spruce, [[Biology:Blue spruce|blue spruce]] (''Picea pungens''), and Norway spruces throughout Ontario, causing severe defoliation and sometimes killing small, stressed trees. White spruce is intermediately susceptible. Dead needles show rows of black fruiting bodies. Infection usually begins on lower branches. On white spruce, infected needles are usually retained on the tree into the following summer. The fungicide Chlorthalonil is registered for controlling this needle cast (Davis 1997).<ref name="davis">Davis, C. (24 September 1997) "Tree talk". ''The Sault Star''. Marie, Ontario. p. B2.</ref>

==== ''Valsa kunzei'' branch and stem canker ====
A branch and stem canker associated with the [[Biology:Fungus|fungus]] ''Valsa kunzei'' Fr. var. ''picea'' was reported on white and Norway spruces in Ontario (Jorgensen and Cafley 1961)<ref name="jorg">{{cite journal |author=Jorgensen, E. and Cafley, J.D. |year=1961 |title=Branch and stem cankers of white and Norway spruces in Ontario |journal=For. Chron. |volume=37 |issue=4 |pages=394–400 |doi=10.5558/tfc37394-4|doi-access=free }}</ref> and Quebec (Ouellette and Bard 1962).<ref name="ouel">{{cite journal |author=Ouellette, G.B.; Bard, G. |year=1962 |title=Observations on a canker and resinosis in white and Norway spruce |journal=Can. Dep. For., for. Ent. Path. Branch, Ottawa ON, Bi-mo. Progr. Rep. |volume=18 |issue=2 |pages=2}}</ref> In Ontario, only trees of low vigour were affected, but in Quebec vigorous trees were also infected.

=== Predators ===
[[File:Spruce in the park of Peterhof.jpg|thumb|Spruce in the park of Peterhof]]
Small [[Biology:Mammal|mammal]]s ingest [[Biology:Pinophyta|conifer]] seeds, and also consume [[Biology:Seedling|seedling]]s. Cage feeding of deer mice (''Peromyscus maniculatus'') and red-backed vole (''Myodes gapperi'') showed a daily maximum seed consumption of 2000 white spruce seeds and of 1000 seeds of [[Biology:Pinus contorta|lodgepole pine]], with the 2 species of mice consuming equal amounts of seed, but showing a preference for the pine over the spruce (Wagg 1963).<ref name="wagg1">{{cite journal |author=Wagg, J.W.B. |year=1963 |title=Notes on food habits of small mammals of the white spruce forest |journal=For. Chron. |volume=39 |issue=4 |pages=436–445 |doi=10.5558/tfc39436-4 |doi-access=free}}</ref> The short-tailed meadow vole (''Microtus pennsylvanicus'' Ord) voraciously ate all available white spruce and lodgepole pine seedlings, pulling them out of the ground and holding them between their front feet until the whole seedling had been consumed. Wagg (1963)<ref name="wagg1" /> attributed damage observed to the bark and cambium at ground level of small white spruce seedlings over several seasons to meadow voles.

Once shed, seeds contribute to the diet of small mammals, e.g., [[Biology:Peromyscus|deer mice]], [[Biology:Clethrionomys|red-backed vole]]s, [[Biology:Montane vole|mountain voles]] (''Microtus montanus''), and [[Biology:Chipmunk|chipmunk]]s (''Eutamias minimus''). The magnitude of the loss is difficult to determine, and studies with and without seed protection have yielded conflicting results. In western Montana, for example, spruce seedling success was little better on protected than on unprotected seed spots (Schopmeyer and Helmers 1947),<ref name="schop">Schopmeyer, C.S.; Helmers, A.E. 1947. Seeding as a means of [[Biology:Reforestation|reforestation]] in the northern Rocky Mountain Region. USDA For. Serv., Washington DC, Circular 772. 30 p.</ref> but in British Columbia spruce regeneration depended on protection from rodents (Smith 1955).<ref name="smith">Smith, J.H.G. 1955 [1956 acc to E3999 bib]. Some factors affecting reproduction of Engelmann spruce and alpine fir. British Columbia Dep. Lands For., For. Serv., Victoria BC, Tech. Publ. 43 p. [Coates et al. 1994, Nienstaedt and Teich 1972]</ref>

An important albeit indirect biotic constraint on spruce establishment is the depredation of seed by squirrels. As much as 90% of a cone crop has been harvested by red squirrels (Zasada et al. 1978).<ref name="zasada">Zasada, J.C.; Foote, M.J.; Deneke, F.J.; Parkerson, R.H. 1978. Case history of an excellent white spruce cone and seed crop in interior Alaska: cone and seed production, germination and seedling survival. USDA, For. Serv., Pacific NW For. Range Exp. Sta., Portland OR, Gen. Tech. Rep. PNW-65. 53 p.</ref> Deer mice, voles, chipmunks, and shrews can consume large quantities of seed; 1 mouse can eat 2000 seeds per night.<ref name="rad2">{{cite journal |last1=Radvanyi |first1=A |year=1970 |title=Small mammals and regeneration of white spruce forests in western Alberta |journal=Ecology |volume=51 |issue=6 |pages=1102–1105 |doi=10.2307/1933641 |jstor=1933641|bibcode=1970Ecol...51.1102R }}</ref> Repeated applications of half a million seeds/ha failed to produce the 750 trees/ha sought by Northwest Pulp and Power, Ltd., near Hinton, Alberta (Radvanyi 1972),<ref name="rad3">Radvanyi, A. 1972. Small mammals and regeneration of white spruce in western Alberta. p. 21–23 ''in'' McMinn, R.G. (Ed.). White Spruce: Ecology of a Northern Resource. Can. Dep. Environ., Can. For. Serv., Edmonton AB, Inf. Rep. NOR-X-40.</ref> but no doubt left a lot of well-fed small mammals. Foraging by squirrels for winter buds (Rowe 1952)<ref name="rowe1">Rowe, J.S. 1952. Squirrel damage to white spruce. Can. Dep. Resour. Devel., For. Branch, For. Res. Div., Ottawa ON, Silv. Leafl. 61. 2 p.</ref> has not been reported in relation to young plantations, but Wagg (1963)<ref name="wagg1" /> noted that at Hinton AB, red squirrels were observed cutting the lateral and terminal twigs and feeding on the vegetative and flower buds of white spruce.

Red squirrels in Alaska have harvested as much as 90% of a cone crop (Zasada et al. 1978);<ref name="zasada" /> their ''modus operandi'' is to cut off great numbers of cones with great expedition early in the fall, and then "spend the rest of the fall shelling out the seeds". In Manitoba, Rowe (1952)<ref name="rowe1" /> ascribed widespread severing of branch tips 5&nbsp;cm to 10&nbsp;cm long on white spruce ranging "from sapling to veteran size" to squirrels foraging for winter buds, cone failure having excluded the more usual food source. The damage has not been reported in relation to small trees, outplants or otherwise.

[[Biology:Porcupine|Porcupine]]s (''Erethizon dorsatum'' L.) may damage spruce (Nienstaedt 1957),<ref name="nien">Nienstaedt, H. 1957. Silvical characteristics of white spruce (''Picea glauca''). USDA, For. Serv., Lake States For. Exp. Sta., St. Paul MN, Pap. 55. 24 p.</ref> but prefer red pine.<ref name="mcleod">{{cite report |url=https://cfs.nrcan.gc.ca/publications?id=30504 |title=Plantations of the Acadia Forest Experiment Station |last=McLeod |first=J.W. |date=1956 |publisher=Can. Dep. Northern Affairs National Resour., For. Branch, For. Res. Div. |id=Tech. Note 31}}</ref> Bark-stripping of white spruce by black bear (''Euarctos americanus perniger'') is locally important in Alaska (Lutz 1951),<ref name="lutz">{{cite journal |author=Lutz, H.J. |year=1951 |title=Damage to trees by black bears in Alaska |journal=J. For. |volume=49 |pages=522–523}}</ref> but the bark of white spruce is not attacked by field mice (''Microtus pennsylvanicus'' Ord),<ref name="little">{{cite journal |author=Littlefield, E.W.; Schoomaker, W.J.; Cook, D.B. |year=1946 |title=Field mouse damage to coniferous plantations |journal=J. For. |volume=44 |pages=745–749}}</ref> even in years of heavy infestation.

=== Pests ===
The eastern spruce budworm (''[[Biology:Choristoneura fumiferana|Choristoneura fumiferana]]'') is a major pest of spruce trees in forests throughout Canada and the eastern United States.<ref name="Powell">{{Cite book |url=https://books.google.com/books?id=YH7WzKFzX0MC&pg=PP1 |title=Biosystematic Studies of Conifer-Feeding Choristoneura (Lepidoptera Tortricidae) in the Western United States |date=1995 |publisher=University of California Press |isbn=9780520097964 |editor-last=Powell |editor-first=Jerry A.}}</ref> Two of the main host plants are [[Biology:Picea mariana|black spruce]] and white spruce.<ref name="bal2">Balch, R.E.; Webb, F.E.; Morris, R.F. (1954). [https://cfs.nrcan.gc.ca/publications?id=8263 Results of spraying against spruce budworm in New Brunswick]. Can. Dep. Agric., For. Biol. Div., Ottawa ON, Bi-mo. Progr. Rep. 10(1).</ref> Population levels oscillate, sometimes reaching extreme outbreak levels that can cause extreme defoliation of and damage to spruce trees. To reduce destruction, there are multiple methods of control in place, including pesticides.<ref name="Pheromone">{{Cite book |last1=Allison |first1=Jeremy D. |url=https://books.google.com/books?id=1GjgDAAAQBAJ |title=Pheromone Communication in Moths: Evolution, Behavior, and Application |last2=Carde |first2=Ring T. |date=2016 |publisher=Univ of California Press |isbn=9780520964433 |pages=265–271}}</ref>

Horntails, or Wood Wasps, use this tree for egg laying and the larvae will live in the outer inch of the tree under the bark.

Spruce beetles (''[[Biology:Dendroctonus rufipennis|Dendroctonus rufipennis]]'') have destroyed swathes of spruce forest in western North America from Alaska to Wyoming.

== Uses ==

===Timber===
[[File:Wood picea abies.jpg|thumb|upright|''P. abies'' wood]]
Spruce is useful as a building wood, commonly referred to by several different names including North American timber, SPF (spruce, pine, fir) and whitewood (the collective name for spruce wood).<ref>{{Cite web |date=2021-09-07 |title=17 Different Types of Pine Wood |url=https://www.homestratosphere.com/types-of-pine-wood/ |access-date=2022-04-27 |website=Home Stratosphere |language=en-US}}</ref> Spruce wood is used for many purposes, ranging from general construction work and crates to highly specialised uses in wooden aircraft.<ref>{{Cite web |last=LaFalce |first=Ben |date=2019-09-24 |title=Types of Wood: Guide to Choose the Best for Your Furniture |url=https://octaneseating.com/blog/types-of-wood-for-furniture/ |access-date=2022-04-27 |website=Octane Seating |language=en-US}}</ref> The [[Biography:Wright brothers|Wright brothers]]' first aircraft, the ''[[Engineering:Wright Flyer|Flyer]]'', was built of spruce.<ref>{{cite web |url=http://www.nasm.si.edu/exhibitions/gal100/wright1903.html |title=Milestones of Flight - 1903 Wright Flyer |publisher=Smithsonian National Air and Space Museum |url-status=dead |archive-url=https://web.archive.org/web/20040405082131/http://www.nasm.si.edu/exhibitions/gal100/wright1903.html |archive-date=5 April 2004 |access-date=21 August 2016|date=2016-04-28}}</ref>

Because this species has no insect or decay resistance qualities after logging, it is generally recommended for construction purposes as indoor use only (indoor drywall framing, for example). Spruce wood, when left outside cannot be expected to last more than 12–18 months depending on the type of climate it is exposed to.<ref>{{cite web |title=Picea Genus (spruce) |url=https://conifersociety.org/conifers/picea |website=American Conifer Society |access-date=13 June 2019}}</ref>

===Pulpwood===
Spruce is one of the most important woods for paper uses, as it has long wood fibres which bind together to make strong paper. The fibres are thin walled and collapse to thin bands upon drying. Spruces are commonly used in mechanical pulping as they are easily [[Chemistry:Bleaching of wood pulp|bleached]]. Together with northern [[Biology:Pine|pine]]s, northern spruces are commonly used to make NBSK. Spruces are cultivated over vast areas as pulpwood.

===Food and medicine===
[[File:SpruceEssentialOil.png|thumb|Spruce (''Picea mariana'') essential oil in a clear glass vial]]

The fresh shoots of many spruces are a natural source of [[Biology:Vitamin C|vitamin C]].<ref>{{cite web |url=http://www.for.gov.bc.ca/hfd/library/documents/treebook/sitkaspruce.htm |title= Tree Book - Sitka spruce (''Picea sitchensis'') |publisher=British Columbia Ministry of Forests, Lands and Natural Resource Operations |access-date=29 July 2006}}</ref> Captain Cook made alcoholic sugar-based spruce beer during his sea voyages in order to prevent [[Medicine:Scurvy|scurvy]] in his crew.<ref>{{cite book |url=https://books.google.com/books?id=P1Oy7Qz1tewC&q=Spruce+beer+Captain+Cook&pg=PA39 |title=A social history of medicines in the twentieth century: to be taken three times a day |first=J. K. |last=Crellin |page=39 |location=New York |publisher=Pharmaceutical Products Press |year=2004 |isbn=978-0789018441}}</ref><ref>{{cite journal |pmid=12810402 |title=Captain Cook's beer: the antiscorbutic use of malt and beer in late 18th century sea voyages |first=Brett J. |last=Stubbs |journal=Asia Pacific Journal of Clinical Nutrition |volume=12 |issue=2 |date=June 2003 |pages=129–137}}</ref> The leaves and branches, or the essential oils, can be used to brew spruce beer.

In Finland, young spruce [[Biology:Bud|bud]]s are sometimes used as a spice, or boiled with sugar to create spruce bud syrup.<ref>{{Cite web|title=Kadoksissa ollut juomaresepti löytyi – kuusenkerkästä tehdään muutakin kuin siirappia|url=http://yle.fi/uutiset/kadoksissa_ollut_juomaresepti_loytyi__kuusenkerkasta_tehdaan_muutakin_kuin_siirappia/8926774|access-date=2021-08-08|website=Yle Uutiset|date=6 June 2016|language=fi}}</ref><ref>{{cite journal |vauthors=Jyske T, Järvenpää E, Kunnas S, Sarjala T, Raitanen JE, Mäki M, Pastell H, Korpinen R, Kaseva J, Tupasela T |title=Sprouts and Needles of Norway Spruce (''Picea abies'' (L.) Karst.) as Nordic Specialty-Consumer Acceptance, Stability of Nutrients, and Bioactivities during Storage |journal=Molecules |volume=25 |issue=18 |pages=4187 |date=2020 |doi=10.3390/molecules25184187 |pmid=32932686 |pmc=7570650|doi-access=free }}</ref> In survival situations spruce needles can be directly ingested or boiled into a tea. This replaces large amounts of vitamin C. Also, water is stored in a spruce's needles, providing an alternative means of hydration {{Clarify|date=June 2009}}. Spruce can be used as a preventive measure for [[Medicine:Scurvy|scurvy]] in an environment where meat is the only prominent food source {{Clarify|date=June 2009}}.

===Tonewood===
Spruce is the standard material used in soundboards for many musical instruments, including [[Engineering:Guitar|guitar]]s, [[Engineering:Mandolin|mandolin]]s, [[Engineering:Cello|cello]]s, [[Engineering:Violin|violin]]s, and the soundboard at the heart of a [[Engineering:Piano|piano]] and the [[Engineering:Harp|harp]].
Wood used for this purpose is referred to as tonewood.

Spruce, along with [[Biology:Thuja plicata|cedar]], is often used for the soundboard/top of an acoustic guitar. The main types of spruce used for this purpose are Sitka, Engelmann, Adirondack and European spruces.

===Other uses===
The resin was used in the manufacture of [[Chemistry:Pitch (resin)|pitch]] in the past (before the use of [[Chemistry:Petrochemical|petrochemical]]s); the scientific name ''Picea'' derives from [[Social:Latin|Latin]] {{wikt-lang|la|picea}} "pitch pine" (referring to [[Biology:Scots pine|Scots pine]]),<ref>{{L&S|picea|pĭcĕa|ref}}</ref> from {{wikt-lang|la|piceus}}, an adjective from {{wikt-lang|la|pix}} "pitch".

Native Americans in [[Earth:North America|North America]] use the thin, pliable roots of some species for weaving [[Engineering:Basket|basket]]s and for sewing together pieces of [[Biology:Birch|birch]] bark for canoes. See also Kiidk'yaas for an unusual golden Sitka Spruce sacred to the Haida people.

Spruces are popular ornamental trees in [[Earth:Horticulture|horticulture]], admired for their evergreen, symmetrical narrow-conic growth habit. For the same reason, some (particularly ''Picea abies'' and ''P. omorika'') are also extensively used as Christmas trees, with artificial Christmas trees often being produced in their likenesses.

Spruce branches are also used at Aintree racecourse, Liverpool, to build several of the fences on the Grand National course. Spruce wood is also used to make sculptures.

== Genome ==
The nuclear,<ref>{{Cite journal | url=https://www.ncbi.nlm.nih.gov/nuccore/ALWZ000000000.4/ |title = Picea glauca, whole genome shotgun sequencing project|date = 13 March 2015}}</ref> mitochondrial<ref>{{Cite journal | url=https://www.ncbi.nlm.nih.gov/nuccore/LKAM00000000 |title = Picea glauca, whole genome shotgun sequencing project|date = 12 January 2016}}</ref><ref>{{cite journal|pmid=26645680|pmc=4758241 |author=Jackman SD, Warren RL, Gibb EA, Vandervalk BP, Mohamadi H, Chu J, Raymond A, Pleasance S, Coope R, Wildung MR, Ritland CE, Bousquet J, Jones SJM, Bohlmann J, Birol I. |title=Organellar Genomes of White Spruce (''Picea glauca''): Assembly and Annotation|journal=Genome Biology and Evolution |volume=8 |issue=1 |pages=29–41 |year=2015 |doi=10.1093/gbe/evv244}}</ref> and chloroplast<ref>{{Cite journal | url=https://www.ncbi.nlm.nih.gov/nuccore/NC_028594.1 | title=''Picea glauca'' chloroplast, complete genome| date=3 December 2015}}</ref> genomes of British Columbia interior spruce have been sequenced. The large (20 Gbp) nuclear genome and associated gene annotations of interior spruce (genotype PG29) were published in 2013<ref>{{cite journal|author=Birol I, Raymond A, Jackman SD, Pleasance S, Coope R, Taylor GA, Yuen MM, Keeling CI, Brand D, Vandervalk BP, Kirk H, Pandoh P, Moore RA, Zhao Y, Mungall AJ, Jaquish B, Yanchuk A, Ritland C, Boyle B, Bousquet J, Ritland K, Mackay J, Bohlmann J, Jones SJ |title= Assembling the 20 Gb white spruce (''Picea glauca'') genome from whole-genome shotgun sequencing data|journal= Bioinformatics|volume= 29|issue= 12|pages= 1492–1497|doi=10.1093/bioinformatics/btt178|pmid= 23698863|pmc= 3673215|year= 2013}}</ref> and 2015.<ref>{{cite journal|author=Warren RL, Keeling CI, Yuen MMS, Raymond A, Taylor GA, Vandervalk BP, Mohamadi H, Paulino D, Chiu R, Jackman SD, Robertson G, Yang C, Boyle B, Hoffmann M, Weigel D, Nelson DR, Ritland C, Isabel N, Jaquish B, Yanchuk A, Bousquet J, Jones SJM, Mackay J, Birol I, Bohlmann J. |title=Improved white spruce (''Picea glauca'') genome assemblies and annotation of large gene families of conifer terpenoid and phenolic defense metabolism|journal=The Plant Journal|volume=83|issue=2|pages=189–212|doi=10.1111/tpj.12886|pmid=26017574|year=2015|doi-access=free}}</ref>

==References==
{{Reflist}}

==External links==
{{AmCyc Poster}}
*theplantlist.org / [http://www.theplantlist.org/tpl1.1/search?q=picea ''Picea'' (Spruce)]
*conifers.org / [http://www.conifers.org/pi/Picea.php Gymnosperm Database - ''Picea'']
*efloras.org / [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=125375 ''Picea'']
*pinetum.org / Arboretum de Villardebelle: Cones of selected species of ''Picea'': [http://www.pinetum.org/cones/PICEAcones.htm page 1], [http://www.pinetum.org/cones/PICEAcones2.htm Arboretum de Villardebelle page 2]

{{Plant classification}}
{{Taxonbar|from=Q26782}}

[[Category:Picea| ]]

{{Sourceattribution|Spruce}}