Editing Pinaceae (section)

==Classification==
[[File:Ab plant 673.jpg|thumb|An immature second-year cone of [[European black pine]] (''Pinus nigra'') with the light brown umbo visible on the green cone scales]]
[[File:Norway Spruce cone.jpg|thumb|An immature cone of [[Norway spruce]] (''Picea abies'') with no umbo]]

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. An 1891 publication divided the family into two subfamilies, using the number and position of resin canals in the primary vascular region of the young taproot as the primary consideration. In a 1910 publication, the family was divided into two tribes based on the occurrence and type of long–short shoot dimorphism.

A more recent classification divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family. Below is an example of how the morphology has been used to classify Pinaceae. 
The 11 genera are grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves:<ref>{{cite journal |author=Robert A. Price, Jeanine Olsen-Stojkovich & Jerold M. Lowenstein |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}}</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.
* 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.
* 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.
* 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.

=== Phylogeny ===
A revised 2018 phylogeny places ''Cathaya'' as sister to the pines rather than in the Laricoidae subfamily with ''Larix'' and ''Pseudotsuga''.

{| class="wikitable"
|-
! colspan=1 |Ran et al. 2018<ref>{{Cite journal|last1=Ran|first1=Jin-Hua|last2=Shen|first2=Ting-Ting|last3=Wu|first3=Hui|last4=Gong|first4=Xun|last5=Wang|first5=Xiao-Quan|date=2018-12-01|title=Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis|url=http://www.sciencedirect.com/science/article/pii/S1055790318301246|journal=Molecular Phylogenetics and Evolution|language=en|volume=129|pages=106–116|doi=10.1016/j.ympev.2018.08.011|pmid=30153503 |bibcode=2018MolPE.129..106R |s2cid=52110440 |issn=1055-7903|url-access=subscription}}</ref> & Leslie et al. 2018<ref>{{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 |display-authors=et al. |year=2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=American Journal of Botany |url=https://doi.org/10.1002/ajb2.1143 |volume=105 |issue=9 |pages=1531–1544 | doi=10.1002/ajb2.1143 |pmid= 30157290|pmc= |bibcode= |doi-access=}}</ref><ref>{{cite journal |last1=Leslie |first1=Andrew B. |display-authors=et al. |year=2018 |title=ajb21143-sup-0004-AppendixS4 |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |url=https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf |doi-access=|pmid=30157290 }}</ref>
! colspan=1 |Stull et al. 2021<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= 2021NatPl...7.1015S|s2cid=232282918 |doi-access=}}</ref><ref>{{cite web |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>
|- 
| style="vertical-align:top|
{{clade| style=font-size:90%;line-height:80%;
|1={{clade
  |label1=[[Abietoideae]]
  |1={{clade
    |1={{clade
      |label1=Cedreae
      |1=''[[Cedrus]]'' (cedars 4 sp.)
       }}
    |2={{clade
      |label1=Pseudolariceae
      |1={{clade
        |1=''[[Pseudolarix]]'' (golden larch 1 sp.)
        |2={{clade
          |1=''[[Nothotsuga]]'' (1 sp.)
          |2=''[[Tsuga]]'' (hemlock  9 sp.)
           }}
         }}
      |label2=Abieteae
      |2={{clade
        |1=''[[Keteleeria]]'' (3 sp.)
        |2=''[[Abies]]'' (firs c.50 sp.)
         }}
       }}
     }}
  |label2=[[Pinoideae]]
  |2={{clade
    |label1=Lariceae
    |1={{clade
      |1=''[[Pseudotsuga]]'' (Douglas-firs 5 sp.)
      |2=''[[Larix]]'' (larches 14 sp.)
       }}
    |label2=Pineae
    |2={{clade
      |1=''[[Picea]]'' (spruces c 35 sp.)
      |2={{clade
        |1=''[[Cathaya]]'' (1 sp.)
        |2=''[[Pinus]]'' (pines c.115 sp.)
         }}
       }}
     }}
   }}
}}
|
{{clade| style=font-size:90%;line-height:80%;
|1={{clade
  |label1=[[Abietoideae]]
  |1={{clade
    |1={{clade
      |label1=Cedreae
      |1=''[[Cedrus]]''
       }}
    |2={{clade
      |label1=Pseudolariceae
      |1={{clade
        |1=''[[Pseudolarix]]''
        |2={{clade
          |1=''[[Nothotsuga]]''
          |2=''[[Tsuga]]''
           }}
         }}
      |label2=Abieteae
      |2={{clade
        |1=''[[Keteleeria]]''
        |2=''[[Abies]]''
         }}
       }}
     }}
  |label2=[[Pinoideae]]
  |2={{clade
    |label1=Lariceae
    |1={{clade
      |1=''[[Pseudotsuga]]''
      |2=''[[Larix]]''
       }}
    |label2=Pineae
    |2={{clade
      |1={{clade
        |1=''[[Cathaya]]''
        |2=''[[Picea]]''
         }}
      |2=''[[Pinus]]''
       }}
     }}
   }}
}}
|}

Multiple molecular studies indicate that in contrast to previous classifications placing it outside the conifers, [[Gnetophyta]] may in fact be the sister group to the Pinaceae, with both lineages having diverged during the early-mid [[Carboniferous]]. This is known as the "gnepine" hypothesis.<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|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|language=en|volume=7|issue=8|pages=1015–1025|doi=10.1038/s41477-021-00964-4|pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |issn=2055-0278}}</ref><ref>{{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|pages=20181012|doi=10.1098/rspb.2018.1012|pmc=6030518|pmid=29925623}}</ref>

=== Evolutionary history ===
Pinaceae is estimated to have 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 |language=en |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430 |issn=1537-2197}}</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}}</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 |language=en |volume=99 |issue=4 |pages=708–720 |doi=10.3732/ajb.1100595 |pmid=22491001}}</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 |language=en |volume=47 |pages=379–389 |s2cid=54653621 }}</ref><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><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 |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 cycles.<ref>{{Cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy M. |last3=Rai |first3=Hardeep S. |last4=Crane |first4=Peter R. |last5=Donoghue |first5=Michael J. |last6=Mathews |first6=Sarah |date=2012-10-02 |title=Hemisphere-scale differences in conifer evolutionary dynamics |journal=Proceedings of the National Academy of Sciences |language=en |volume=109 |issue=40 |pages=16217–16221 |doi=10.1073/pnas.1213621109 |doi-access=free |issn=0027-8424 |pmc=3479534 |pmid=22988083|bibcode=2012PNAS..10916217L }}</ref>