View source for Pinaceae

{{Short description|Family of conifers}}
{{Automatic taxobox
| fossil_range = {{fossil range|Jurassic|Recent}} Possible Triassic record <ref>{{cite web |title=Fort Wingate (Triassic of the United States) |url=https://paleobiodb.org/classic/basicCollectionSearch?collection_no=23968 |website=PBDB.org}}</ref>
| image = Glacier-Lyman-Tamarack.jpg
| image_caption = ''Larix'' (golden), ''Abies'' (central foreground) and ''Pinus'' (right foreground)
| taxon = Pinaceae
| authority = [[John Lindley|Lindley]] 1836
| subdivision_ranks = Genera
| subdivision = * '''[[Abietoideae]]'''
** ''[[Abies]]''
** ''[[Cedrus]]''
** ''[[Keteleeria]]''
** ''[[Nothotsuga]]''
** ''[[Pseudolarix]]''
** ''[[Tsuga]]''
* '''[[Pinoideae]]'''
** ''[[Pinus]]''
* '''[[Piceoideae]]'''
** ''[[Picea]]''
* '''[[Laricoideae]]'''
** ''[[Pseudotsuga]]''
** ''[[Cathaya]]''
** ''[[Larix]]''
| synonyms = * Abietaceae <small>von Berchtold & Presl 1820</small>
* Cedraceae <small>Vest 1818</small>
* Compsostrobaceae <small>Delevoryas & Hope 1973</small>
* †Kranneraceae <small>Corda 1866</small>
* Piceaceae <small>Goroschankin 1904</small>
}}

The '''Pinaceae''', or '''pine family''', are [[conifer]] trees or shrubs, including many of the well-known conifers of commercial importance such as [[Cedrus|cedars]], [[fir]]s, [[Tsuga|hemlocks]], [[larch]]es, [[pine]]s and [[spruce]]s. The family is included in the order [[Pinales]], formerly known as [[Coniferales]]. Pinaceae have distinctive cones with woody scales bearing typically two [[Ovule|ovules]], and are supported as [[monophyletic]] by both [[Morphology (biology)|morphological]] trait and genetic analysis.<ref>{{Cite journal |last1=Gernandt |first1=David S. |last2=Holman |first2=Garth |last3=Campbell |first3=Christopher |last4=Parks |first4=Matthew |last5=Mathews |first5=Sarah |last6=Raubeson |first6=Linda A. |last7=Liston |first7=Aaron |last8=Stockey |first8=Ruth A. |last9=Rothwell |first9=Gar W. |date=September 2016 |title=Phylogenetics of extant and fossil Pinaceae: methods for increasing topological stability |url=http://www.nrcresearchpress.com/doi/10.1139/cjb-2016-0064 |journal=Botany |language=en |volume=94 |issue=9 |pages=863–884 |doi=10.1139/cjb-2016-0064 |issn=1916-2790|url-access=subscription }}</ref> They are the largest extant conifer family in species diversity, with between 220 and 250 species (depending on [[Taxonomy (biology)|taxonomic]] opinion) in 11 genera,<ref name="Farjon">{{cite book |author=Aljos Farjon |year=1998 |title=World Checklist and Bibliography of Conifers |publisher=[[Royal Botanic Gardens, Kew]]  |isbn=978-1-900347-54-9}}</ref> and the second-largest (after [[Cupressaceae]]) in geographical range, found in most of the [[Northern Hemisphere]], with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of [[Boreal forest|boreal]], coastal, and [[montane forest]]s. One species, ''[[Pinus merkusii]]'', grows just south of the [[equator]] in Southeast Asia.<ref>{{Gymnosperm Database |family=Pinaceae |genus=Pinus |species=merkusii |access-date=March 17, 2015}}</ref> Major [[centre of diversity|centres of diversity]] are found in the mountains of [[southwest China]], Mexico, central Japan, and [[California]].

==Description==

<!--All-->The members of the family Pinaceae are [[tree]]s (rarely [[shrub]]s) growing from {{convert|2|to|100|m|ft|sigfig=1|abbr=off}} tall, mostly [[evergreen]] (except the [[deciduous]] ''[[Larix]]'' and ''[[Pseudolarix]]''), [[resin]]ous, [[monoecious]], with subopposite or whorled branches, and spirally arranged, linear (needle-like) leaves.<ref name="Farjon"/> The embryos of Pinaceae have three to 24 [[cotyledon]]s.

The female [[conifer cone|cones]] are large and usually woody, {{convert|2|-|60|cm|0|abbr=off}} long, with numerous spirally arranged scales, and two winged [[seed]]s on each scale. The male cones are small, {{convert|0.5|-|6|cm|abbr=on|sigfig=1|frac=4}} long, and fall soon after pollination; pollen dispersal is by wind. Seed dispersal is mostly by wind, but some species have large seeds with reduced wings, and are dispersed by birds. Analysis of Pinaceae cones reveals how selective pressure has shaped the evolution of variable cone size and function throughout the family. Variation in cone size in the family has likely resulted from the variation of seed dispersal mechanisms available in their environments over time. All Pinaceae with seeds weighing less than 90&nbsp;milligrams are seemingly adapted for wind dispersal. Pines having seeds larger than 100&nbsp;mg are more likely to have benefited from adaptations that promote animal dispersal, particularly by birds. Pinaceae that persist in areas where [[tree squirrel]]s are abundant do not seem to have evolved adaptations for bird dispersal.<ref name="Benkman 1995">{{cite journal |author=Craig W. Benkman |year=1995 |title=Wind dispersal capacity of pine seeds and the evolution of different seed dispersal modes in pines |journal=[[Oikos (journal)|Oikos]] |volume=73 |issue=2 |pages=221–224 |jstor=3545911 |url=http://www.uwyo.edu/benkman/pdfs%20of%20papers/benkman_oikos_1995.pdf |doi=10.2307/3545911|bibcode=1995Oikos..73..221B }}</ref>

[[wikt:Boreal|Boreal]] conifers have multiple [[adaptation]]s to survive winters, including the tree's conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss.<ref name="Michigan">{{cite web |title=Winter Adaptations of Trees |url=https://mff.forest.mtu.edu/Environment/WinterTrees.htm |publisher=[[Michigan Technological University]] |access-date=20 September 2025}}</ref>

<gallery class=center mode=nolines widths=200 heights=260>
File:496 Pinus silvestris.jpg|Features of ''[[Pinus sylvestris]]''
File:Vagamon Pine Forest.jpg|Cultivated pine forest in [[Western Ghats]], India
</gallery>

== Evolution ==

=== Fossil history ===

The Pinaceae diverged from other conifer groups during the late [[Carboniferous]] ~313 million years ago.<ref name="Leslie 2018 1531–1544">{{Cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |date=2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |url=https://bsapubs.onlinelibrary.wiley.com/doi/abs/10.1002/ajb2.1143 |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430 |url-access=subscription }}</ref> Various possible [[stem-group]] relatives have been reported from as early as the Late [[Permian]] ([[Lopingian]]) The extinct conifer cone genus ''[[Schizolepidopsis]]'' likely represent stem-group members of the Pinaceae, the first good records of which are in the Middle-Late [[Triassic]], with abundant records during the [[Jurassic]] across Eurasia.<ref>{{Cite journal |last1=Domogatskaya |first1=Ksenia V. |last2=Herman |first2=Alexei B. |date=May 2019 |title=New species of the genus Schizolepidopsis (conifers) from the Albian of the Russian high Arctic and geological history of the genus |url=https://linkinghub.elsevier.com/retrieve/pii/S0195667118304257 |journal=Cretaceous Research |language=en |volume=97 |pages=73–93 |doi=10.1016/j.cretres.2019.01.012|bibcode=2019CrRes..97...73D |s2cid=134849082 |url-access=subscription }}</ref><ref name=":11">{{Cite journal |last1=Matsunaga |first1=Kelly K. S. |last2=Herendeen |first2=Patrick S. |last3=Herrera |first3=Fabiany |last4=Ichinnorov |first4=Niiden |last5=Crane |first5=Peter R. |last6=Shi |first6=Gongle |date=2021-05-10 |title=Ovulate Cones of Schizolepidopsis ediae sp. nov. Provide Insights into the Evolution of Pinaceae |journal=International Journal of Plant Sciences |volume=182 |issue=6 |pages=490–507 |doi=10.1086/714281 |issn=1058-5893 |doi-access=free|bibcode=2021IJPlS.182..490M }}</ref> The oldest [[crown group]] (descendant of the last common ancestor of all living species) member of Pinaceae is the cone ''[[Eathiestrobus]]'', known from the Upper Jurassic (lower [[Kimmeridgian]], 157.3-154.7 million years ago) of Scotland,<ref>{{Cite journal |last1=Rothwell |first1=Gar W. |last2=Mapes |first2=Gene |last3=Stockey |first3=Ruth A. |last4=Hilton |first4=Jason |date=April 2012 |title=The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae |journal=American Journal of Botany |volume=99 |issue=4 |pages=708–720 |doi=10.3732/ajb.1100595 |pmid=22491001|bibcode=2012AmJB...99..708R }}</ref> which likely belongs to the pinoid grouping of the family.<ref name=":12">{{Cite journal |last1=Smith |first1=Selena Y. |last2=Stockey |first2=Ruth A. |last3=Rothwell |first3=Gar W. |last4=Little |first4=Stefan A. |date=2017-01-02 |title=A new species of Pityostrobus (Pinaceae) from the Cretaceous of California: moving towards understanding the Cretaceous radiation of Pinaceae |url=https://www.tandfonline.com/doi/full/10.1080/14772019.2016.1143885 |journal=Journal of Systematic Palaeontology |language=en |volume=15 |issue=1 |pages=69–81 |doi=10.1080/14772019.2016.1143885 |bibcode=2017JSPal..15...69S |s2cid=88292891 |issn=1477-2019|url-access=subscription }}</ref><ref name=":11" /> Pinaceae rapidly radiated during the [[Early Cretaceous]].<ref name="Leslie 2018 1531–1544" /> Members of the modern genera ''Pinus'' (pines), ''Picea'' (spruce) and ''Cedrus'' (cedar) first appear during the Early Cretaceous.<ref>{{Cite journal |last1=Blokhina |first1=N. I. |last2=Afonin |first2=M. |date=2007 |title=Fossil wood Cedrus penzhinaensis sp. nov. (Pinaceae) from the Lower Cretaceous of north-western Kamchatka (Russia) |journal=Acta Paleobotanica |volume=47 |pages=379–389 |s2cid=54653621 }}</ref><ref>{{cite journal |author1=Ashley A. Klymiuk |author2=Ruth A. Stockey |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|bibcode=2012AmJB...99.1069K }}</ref><ref>{{cite journal |author1=Patricia E. Ryberg |author2=Gar W. Rothwell |author3=Ruth A. Stockey |author4=Jason Hilton |author5=Gene Mapes |author6=James B. Riding |year=2012 |title=Reconsidering Relationships among Stem and Crown Group Pinaceae: Oldest Record of the Genus ''Pinus'' from the Early Cretaceous of Yorkshire, United Kingdom |journal=International Journal of Plant Sciences |volume=173 |issue=8 |pages=917–932 |doi=10.1086/667228 |bibcode=2012IJPlS.173..917R |s2cid=85402168}}</ref> The extinct Cretaceous genera ''[[Pseudoaraucaria]]'' and ''[[Obirastrobus]]'' appear to be members of Abietoideae, while ''[[Pityostrobus]]'' appears to be non-monophyletic, containing many disparately related members of Pinaceae.<ref name=":12" /> While Pinaceae, and indeed all of its subfamilies, substantially predate the break up of the super-continent [[Pangaea|Pangea]], its distribution was limited to northern [[Laurasia]].  During the Cenozoic, Pinaceae had higher rates of species turnover than Southern Hemisphere conifers, thought to be driven by range shifts in response to glacial cycle.

=== Phylogeny ===

Molecular studies show that [[Gnetophyta]] is the sister group to the Pinaceae, the lineages having diverged during the early-mid [[Carboniferous]]. This is known as the "gnepine" hypothesis.<ref name=Stull>{{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 |display-authors=5|date=July 19, 2021|title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms|url=https://www.nature.com/articles/s41477-021-00964-4 |journal=Nature Plants |volume=7|issue=8|pages=1015–1025|doi=10.1038/s41477-021-00964-4 |pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |url-access=subscription}}</ref><ref  name="Ran 2018">{{Cite journal |last1=Ran|first1=Jin-Hua|last2=Shen|first2=Ting-Ting|last3=Wang|first3=Ming-Ming|last4=Wang|first4=Xiao-Quan|title=Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms|journal=Proceedings of the Royal Society B: Biological Sciences|year=2018 |volume=285|issue=1881|article-number=20181012|doi=10.1098/rspb.2018.1012|pmc=6030518|pmid=29925623}}</ref><ref name="Liston">{{cite conference |last1=Gernandt |first1=D.S. |last2=Vining |first2=T.F. |last3=Campbell |first3=C.S. |last4=Piñero |first4=D. |last5=Liston |first5=A. |date=August 1999 |title=Molecular phylogeny of Pinaceae and Pinus |conference=IV International Conifer Conference 615 |pages=107-114 |url=https://www.researchgate.net/profile/Aaron-Liston/publication/253545994_Molecular_Phylogeny_of_Pinaceae_and_Pinus/links/0c9605371160789a71000000/Molecular-Phylogeny-of-Pinaceae-and-Pinus.pdf}}</ref> 
The Abietoideae and the Pinoideae diverged in the Jurassic. Pineae and Lariceae diverged in the Late Jurassic, while the Abieteae and Pseudolariceae diverged in the Cretaceous.<ref name=Wang>{{cite journal |last1=Wang |first1=X.-Q. |last2=Tank |first2=D. C. |last3=Sang |first3=T. |title=Phylogeny and Divergence Times in Pinaceae: Evidence from Three Genomes |journal=[[Molecular Biology and Evolution]] |issue=17 |pages=773-781 |url=https://academic.oup.com/mbe/article-pdf/17/5/773/23445669/mbev_17_05_0773.pdf |date=2000 }}</ref>

{{clade
|label1='''Pinaceae'''
|1={{clade
   |label1=[[Abietoideae]]
   |1={{clade
      |label1=Cedreae
      |1=''[[Cedrus]]'' [[File:Cedrus BHL677922.jpg|50px]]
      |2={{clade
         |label1=Pseudolariceae
         |1={{clade
            |label1=''[[Pseudolarix]]'' |sublabel1=(golden larches)
            |1=[[File:Flore des serres v17 111a (cropped).jpg|70px]]
            |2={{clade
               |label1=''[[Nothotsuga]]'' |sublabel1=(bristlecone hemlock)
               |1=[[File:Nothotsuga cones and leaves.jpg|70px]]
               |sublabel2=(hemlocks) |label2=''[[Tsuga]]'' 
               |2=[[File:Tsuga cone and leaves.png|50px]]
               }}
            }}
         |label2=Abieteae
         |2={{clade
            |label1=''[[Keteleeria]]'' 
            |1=[[File:Keteleeria cone and leaves.jpg|65px]]
            |sublabel2=(firs) |label2=''[[Abies]]''
            |2=[[File:Abies alba - Köhler–s Medizinal-Pflanzen-001.jpg|50px]]
            }}
         }}
      }}
   |label2=[[Pinoideae]]
   |2={{clade
      |label1=Lariceae
      |1={{clade
         |label1=''[[Pseudotsuga]]'' |sublabel1=(Douglas-firs)
         |1=[[File:Pinetum woburnense (Douglas firs, page 185).jpg|50px]]
         |label2=''[[Larix]]'' |sublabel2=(larches) 
         |2=[[File:395 Larix europaea DC.jpg|60px]]
         }}
      |label2=Pineae
      |2={{clade
         |1={{clade
            |label1=''[[Cathaya]]'' |sublabel1=(1 sp.<!--no common name-->)
            |1=[[File:Cathaya argyrophylla leaves and cones.jpg|50px]]
            |label2=''[[Picea]]'' |sublabel2=(spruces) 
            |2=[[File:Pinetum woburnense (Spruce, page 146) (cropped).jpg|50px]]
            }}
         |label2=''[[Pinus]]'' |sublabel2=(pines)
         |2=[[File:496 Pinus silvestris.jpg|50px]]
         }}
      }}
   }}
}}

A study by J. D. Lockwood and colleagues in 2013 produced broadly similar results, but with different placements for ''Pseudolarix'' and ''Cathaya''. In this scheme, Pseudolariceae is subsumed by Abieteae.<ref name="Lockwood-2013">{{cite journal |last1=Lockwood |first1=Jared D. |last2=Aleksić |first2=Jelena M. |last3=Zou |first3=Jiabin |last4=Wang |first4=Jing |last5=Liu |first5=Jianquan |last6=Renner |first6=Susanne S. |title=A new phylogeny for the genus Picea from plastid, mitochondrial, and nuclear sequences |journal=Molecular Phylogenetics and Evolution |volume=69 |issue=3 |date=2013 |doi=10.1016/j.ympev.2013.07.004 |pages=717–727 |url=https://linkinghub.elsevier.com/retrieve/pii/S1055790313002819 |access-date=2025-11-07}}</ref>

{{clade
|label1='''Pinaceae'''
|1={{clade
   |label1=[[Abietoideae]]
   |1={{clade
      |label1=Cedreae
      |1=''[[Cedrus]]'' [[File:Cedrus BHL677922.jpg|50px]]
      |label2=Abieteae
      |2={{clade
         |1={{clade
            |label1=''[[Pseudolarix]]'' |sublabel1=(golden larches)
            |1=[[File:Flore des serres v17 111a (cropped).jpg|70px]]
            |2={{clade
               |1={{clade
                  |label1=''[[Nothotsuga]]'' |sublabel1=(bristlecone hemlock)
                  |1=[[File:Nothotsuga cones and leaves.jpg|70px]]
                  |sublabel2=(hemlocks) |label2=''[[Tsuga]]'' 
                  |2=[[File:Tsuga cone and leaves.png|50px]]
                  }}
               |2={{clade
                  |label1=''[[Keteleeria]]'' 
                  |1=[[File:Keteleeria cone and leaves.jpg|65px]]
                  |sublabel2=(firs) |label2=''[[Abies]]''
                  |2=[[File:Abies alba - Köhler–s Medizinal-Pflanzen-001.jpg|50px]]
                  }}
               }}
            }}
         }}
      }}
   |label2=[[Pinoideae]]
   |2={{clade
      |label1=Lariceae
      |1={{clade
         |label1=''[[Pseudotsuga]]'' |sublabel1=(Douglas-firs)
         |1=[[File:Pinetum woburnense (Douglas firs, page 185).jpg|50px]]
         |label2=''[[Larix]]'' |sublabel2=(larches) 
         |2=[[File:395 Larix europaea DC.jpg|60px]]
         }}
      |label2=Pineae
      |2={{clade
         |1={{clade
            |label1=''[[Pinus]]'' |sublabel1=(pines)
            |1=[[File:496 Pinus silvestris.jpg|50px]]
            |label2=''[[Picea]]'' |sublabel2=(spruces) 
            |2=[[File:Pinetum woburnense (Spruce, page 146) (cropped).jpg|50px]]
            }}
         |label2=''[[Cathaya]]'' |sublabel2=(1 sp.)
         |2=[[File:Cathaya argyrophylla leaves and cones.jpg|50px]]
         }}
      }}
   }}
}}

=== Taxonomic history ===

Classification of the subfamilies and genera of Pinaceae has been subject to debate in the past. Pinaceae ecology, morphology, and history have all been used as the basis for methods of analyses of the family. In 1891, [[Philippe Édouard Léon Van Tieghem|Van Tieghem]] divided the family into two subfamilies, using the number and position of [[resin canal]]s in the primary vascular region of the young taproot as the primary consideration. In 1910, [[Friedrich Karl Max Vierhapper |Friedrich Vierhapper]] divided the family into two tribes based on the occurrence and type of long–short shoot dimorphism.<ref>{{cite book |last=Vierhapper |first=Friedrich |author-link=Friedrich Karl Max Vierhapper |title=Entwurf eines neuen Systemes der Coniferen |language=de |trans-title=Proposal for a New System for the Conifers |volume=5 |issue=4 |publisher=G. Fischer |year=1910}}</ref> In 1976, Charles Miller divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family.<ref name="Miller 1976">{{cite journal |last=Miller |first=Charles N. |title=Early evolution in the Pinaceae |journal=Review of Palaeobotany and Palynology |volume=21 |issue=1 |date=1976 |doi=10.1016/0034-6667(76)90024-5 |pages=101–117}}</ref> 

<gallery class=center mode=nolines widths=180 heights=180 caption="Cone features used in Pinaceae taxonomy">
File:Ab plant 673.jpg|Immature 2nd-year cone of ''Pinus nigra'', light brown umbo on green cone scales
File:Norway Spruce cone.jpg|Immature cone of ''Picea abies'', no umbo
</gallery>

For example, Price (1987) classified the Pinaceae into 11 genera, grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves:<ref name="Price 1987">{{cite journal |author=Price, Robert A.; Olsen-Stojkovich, Jeanine; Lowenstein, Jerold M. |year=1987 |title=Relationships among the genera of Pinaceae: an immunological comparison |journal=[[Systematic Botany]] |volume=12 |issue=1 |pages=91–97 |jstor=2419217 |doi=10.2307/2419217 |bibcode=1987SysBo..12...91P }}</ref>

* Subfamily [[Pinoideae]] (''[[Pinus]]''): cones are biennial, rarely triennial, with each year's scale-growth distinct, forming an umbo on each scale, the cone scale base is broad, concealing the seeds fully from [[abaxial]] (below the [[phloem]] vessels) view, the seed is without resin vesicles, the seed wing holds the seed in a pair of claws, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.<ref name="Price 1987"/>
* Subfamily [[Piceoideae]] (''[[Picea]]''): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, seed is without resin vesicles, blackish, the seed wing holds the seed loosely in a cup, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.<ref name="Price 1987"/>
* Subfamily [[Laricoideae]] (''[[Larix]]'', ''[[Pseudotsuga]]'', and ''[[Cathaya]]''): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, the seed is without resin vesicles, whitish, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial only.<ref name="Price 1987"/>
* Subfamily [[Abietoideae]] (''[[Abies]]'', ''[[Cedrus]]'', ''[[Pseudolarix]]'', ''[[Keteleeria]]'', ''[[Nothotsuga]]'', and ''[[Tsuga]]''): cones are annual, without a distinct umbo, the cone scale base is narrow, with the seeds partly visible in abaxial view, the seed has resin vesicles, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial only.<ref name="Price 1987"/>

== Genera ==

=== Extant ===

* ''[[Abies]]'' 
* ''[[Cedrus]]'' 
* ''[[Keteleeria]]'' 
* ''[[Nothotsuga]]''  
* ''[[Pseudolarix]]'' 
* ''[[Tsuga]]'' 
* ''[[Pinus]]'' 
* ''[[Picea]]'' 
* ''[[Pseudotsuga]]'' 
* ''[[Cathaya]]''
* [[Larch|''Larix'']]

=== Extinct ===

* †''[[Tsugaepollenites]]''
* †''[[Abietipites]]''
* †''[[Cedripites]]''
* †''[[Schizolepidopsis]]''
* †''[[Eathiestrobus]]''
* †''[[Pesudoaraucaria]]''
* †''[[Obirostrobus]]''
* †''[[Pityostrobus]]''

== Defense mechanisms ==

{{see also|Plant defense against herbivory}}

External stresses on plants have the ability to change the structure and composition of [[Forest ecology |forest ecosystems]]. Common external stresses that Pinaceae experience are [[herbivore]] and [[pathogen]] attacks, which can kill trees.<ref name=":7">{{Cite journal |last1=Cherubini |first1=Paolo |last2=Fontana |first2=Giovanni |last3=Rigling |first3=Daniel |last4=Dobbertin |first4=Matthias |last5=Brang |first5=Peter |last6=Innes |first6=John L. |date=2002 |title=Tree-Life History Prior to Death: Two Fungal Root Pathogens Affect Tree-Ring Growth Differently |jstor=3072253 |journal=Journal of Ecology |volume=90 |issue=5 |pages=839–850 |doi=10.1046/j.1365-2745.2002.00715.x |doi-access=free |bibcode=2002JEcol..90..839C }}</ref> In order to combat these stresses, trees need to adapt or evolve defenses against these stresses. Pinaceae have evolved myriad mechanical and chemical defenses, or a combination of the two, in order to protect themselves against antagonists.<ref name=":1">{{Cite journal |title=Terpenoid biosynthesis and specialized vascular cells of conifer defense. - Semantic Scholar |year=2010 |doi=10.1111/j.1744-7909.2010.00910.x |pmid=20074143 |s2cid=26043965 |last1=Zulak |first1=K. G. |last2=Bohlmann |first2=J. |journal=Journal of Integrative Plant Biology |volume=52 |issue=1 |pages=86–97 |doi-access=free}}</ref> Pinaceae have the ability to up-regulate a combination of constitutive mechanical and [[Chemical defense |chemical strategies]] to further their defenses.<ref name=":8">{{Cite journal |last1=Franceschi |first1=Vincent R. |last2=Krokene |first2=Paal |last3=Christiansen |first3=Erik |last4=Krekling |first4=Trygve |date=2005-08-01 |title=Anatomical and chemical defenses of conifer bark against bark beetles and other pests |journal=New Phytologist |volume=167 |issue=2 |pages=353–376 |doi=10.1111/j.1469-8137.2005.01436.x |pmid=15998390 |doi-access=free |bibcode=2005NewPh.167..353F }}</ref>

Pinaceae defenses are prevalent in the bark of the trees. This part of the tree contributes a complex defensive boundary against external antagonists.<ref name=":0">Franceschi, V. R., P. Krokene, T. Krekling, and E. Christiansen. 2000. Phloem parenchyma cells are involved in local and distance defense response to fungal inoculation or bark-beetle attack in Norway spruce (Pinaceae). American Journal of Botany 87:314-326.</ref> [[Constitutive defense |Constitutive]] and [[Inducible plant defenses against herbivory |induced defenses]] are both found in the bark.<ref name=":0"/><ref name=":6">{{Cite journal |last1=Hudgins |first1=J. W. |last2=Christiansen |first2=E. |last3=Franceschi |first3=V. R. |date=2004-03-01 |title=Induction of anatomically based defense responses in stems of diverse conifers by methyl jasmonate: a phylogenetic perspective |journal=Tree Physiology |language=en |volume=24 |issue=3 |pages=251–264 |doi=10.1093/treephys/24.3.251 |pmid=14704135 |issn=0829-318X |doi-access=free}}</ref><ref name=":9">{{Cite journal |last1=Krokene |first1=P. |last2=Nagy |first2=N. E. |last3=Solheim |first3=H. |date=2008-01-01 |title=Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection |journal=Tree Physiology |volume=28 |issue=1 |pages=29–35 |doi=10.1093/treephys/28.1.29 |pmid=17938111 |issn=0829-318X |doi-access=free}}</ref>

=== Constitutive defenses ===

[[Plant defense against herbivory#Types|Constitutive defense]]s are typically the first line of defenses used against antagonists. These defenses include sclerified cells, lignified periderm cells, and secondary compounds such as [[Phenols |phenolics]] and resins.<ref name=":2">{{Cite journal |last=Sampedro |first=L. |date=2014-09-01 |title=Physiological trade-offs in the complexity of pine tree defensive chemistry |journal=Tree Physiology |volume=34 |issue=9 |pages=915–918 |doi=10.1093/treephys/tpu082 |pmid=25261122 |issn=0829-318X |doi-access=free |hdl=10261/105595 |hdl-access=free}}</ref><ref name=":0"/><ref name=":6"/> Constitutive defenses are always expressed and offer immediate protection from invaders but can be defeated by antagonists that have evolved adaptations to these defense mechanisms.<ref name=":2"/><ref name=":0"/> Common secondary compounds used by Pinaceae are phenolics or polyphenols. These are preserved in [[vacuole]]s of polyphenolic [[Parenchyma |parenchyma cells]] (PP) in the [[Phloem |secondary phloem]].<ref name=":3">{{Cite journal |last1=Nagy |first1=N. E. |last2=Krokene |first2=P. |last3=Solheim |first3=H. |date=2006-02-01 |title=Anatomical-based defense responses of Scots pine (Pinus sylvestris) stems to two fungal pathogens |journal=Tree Physiology |volume=26 |issue=2 |pages=159–167 |doi=10.1093/treephys/26.2.159 |pmid=16356912 |issn=0829-318X |doi-access=free}}</ref><ref name=":9"/>

=== Induced defenses ===

[[Inducible plant defenses against herbivory |Induced defense]] responses need to be activated by certain cues, such as herbivore damage or other biotic signals.<ref name=":2"/>

A common induced defense mechanism used by Pinaceae is resins.<ref name=":4">{{Cite journal |last1=Nagy |first1=Nina E. |last2=Franceschi |first2=Vincent R. |last3=Solheim |first3=Halvor |last4=Krekling |first4=Trygve |last5=Christiansen |first5=Erik |date=2000-03-01 |title=Wound-induced traumatic resin duct development in stems of Norway spruce (Pinaceae): anatomy and cytochemical traits |journal=American Journal of Botany |language=en |volume=87 |issue=3 |pages=302–313 |doi=10.2307/2656626 |jstor=2656626 |pmid=10718991}}</ref> Resins are also one of the primary defenses used against attack.<ref name=":1"/> Resins are short term defenses that are composed of a complex combination of volatile [[Monoterpene |mono]]- (C<sub>10</sub>) and [[sesquiterpene]]s (C<sub>15</sub>) and nonvolatile [[diterpene]] resin acids (C<sub>20</sub>).<ref name=":1"/><ref name=":4"/> They are produced and stored in specialized secretory areas known as resin ducts, resin blisters, or resin cavities.<ref name=":4"/> Resins have the ability to wash away, trap, fend off antagonists, and are also involved in wound sealing.<ref name=":3"/> They are an effective defense mechanism because they have toxic and inhibitory effects on invaders, such as insects or pathogens.<ref name=":5">{{Cite journal |last1=Lewinsohn |first1=Efraim |last2=Gijzen |first2=Mark |last3=Croteau |first3=Rodney |date=1991-05-01 |title=Defense Mechanisms of Conifers: Differences in Constitutive and Wound-Induced Monoterpene Biosynthesis Among Species |journal=Plant Physiology |volume=96 |issue=1 |pages=44–49 |doi=10.1104/pp.96.1.44 |pmid=16668184 |pmc=1080711}}</ref> Resins could have developed as an evolutionary defense against [[bark beetle]] attacks.<ref name=":4"/> One well researched resin present in Pinaceae is [[oleoresin]]. Oleoresin had been found to be a valuable part of the [[Pinophyta |conifer]] defense mechanism against [[Biotic stress |biotic attacks]]. They are found in [[Plant secretory tissue |secretory tissues]] in tree stems, roots, and leaves.<ref name=":5"/>

Many studies use [[methyl jasmonate]] as an antagonist.<ref name=":6"/><ref name=":9"/><ref name=":10">{{Cite journal |last1=Fäldt |first1=Jenny |last2=Martin |first2=Diane |last3=Miller |first3=Barbara |last4=Rawat |first4=Suman |last5=Bohlmann |first5=Jörg |date=2003-01-01 |title=Traumatic resin defense in Norway spruce (Picea abies): Methyl jasmonate-induced terpene synthase gene expression, and cDNA cloning and functional characterization of (+)-3-carene synthase |journal=Plant Molecular Biology |volume=51 |issue=1 |pages=119–133 |doi=10.1023/A:1020714403780 |pmid=12602896 |s2cid=21153303}}</ref> Methyl jasmonate induces defense responses in the stems of multiple Pinaceae species.<ref name=":6"/><ref name=":10"/> Methyl jasmonate stimulates the activation of PP cells and formation of xylem traumatic resin ducts (TD). These are involved in the release of phenolics and resins, both forms of defense mechanism.<ref name=":6"/><ref name=":9"/>

<gallery class=center mode=nolines widths=200 heights=200>
File:Pinceae_Bishop_pine_prickle_cone_pine_pinus_muricata.jpg | Bishop pine cones
File:Pinaceae_Knobcone_Pine_Pinus_attenuata.jpg | Knobcone pine cone
</gallery>

== References ==

{{Reflist}}

==External links==
{{Commons category}}
{{Wikispecies}}

* [http://www.pinetum.org/ Arboretum de Villardebelle] French Arboretum of conifers around the world 
* [http://www.conifers.org/pi/Pinaceae.php Gymnosperm Database] – Pinaceae
* [http://tolweb.org/Pinaceae/21624 Pinaceae on the web page of the Tree-of-Life project]
* [https://www.thespruce.com/pine-trees-from-around-the-world-3269718 40 Pine Trees From Around the World] by The Spruce
* {{Jepson eFlora|32|Pinaceae|link=1|type=key|mode=cs2}}, covers Californian species and much of western North America
* [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10691 Pinaceae in Flora of North America]
* [http://plants.usda.gov/plant-profile?symbol=PINUS Pinus in USDA Plants Database]

{{Acrogymnospermae classification}}
{{Taxonbar|from=Q101680}}
{{Authority control}}
[[Category:Pinaceae| ]]
[[Category:Pinales families]]