Editing Pine (section)

== Evolution ==

=== Taxonomy ===

{{Main list|List of Pinus species |l1=List of ''Pinus'' species}}

Pines are [[gymnosperm]]s. The genus is divided into two subgenera based on the number of [[Vascular bundle|fibrovascular bundles]] in the needle, and the presence or absence of a resin seal on the scales of the mature cones before opening. The subgenera can be distinguished by cone, seed, and leaf characters:
*[[Pinus subg. Pinus|''Pinus'' subg. ''Pinus'']], the yellow, or hard pine group, with cones with a resin seal on the scales, and generally with harder wood; the needle fascicles mostly have a persistent sheath (two exceptions, ''[[Pinus leiophylla]]'' and ''[[Pinus lumholtzii]]'', have deciduous sheaths).<ref name="RHSD"/> The subgenus has also been called ''diploxylon'', on account of its two fibrovascular bundles.<ref name="Mirov">{{cite book |last1=Mirov |first1=N. T. |title=The Genus ''Pinus'' |date=1967 |publisher=Ronald Press |location=New York}}</ref>
*[[Strobus|''Pinus'' subg. ''Strobus'']], [[synonym (biology)|syn.]] ''Pinus'' subg. ''Ducampopinus'', the white, or soft pine, and pinyon pine groups, with cones without a resin seal on the scales, and usually have softer wood; the needle fascicles mostly have a deciduous sheath (one exception, ''[[Pinus nelsonii]]'', has a persistent sheath).<ref name="RHSD"/> The subgenus has also been called ''haploxylon'', on account of its single fibrovascular bundle.<ref name="Mirov"/>

Phylogenetic evidence indicates that both subgenera have a very ancient divergence from one another.<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 |issn=2055-0278 |access-date=January 10, 2022 |archive-date=January 10, 2022 |archive-url=https://web.archive.org/web/20220110174725/https://www.nature.com/articles/s41477-021-00964-4/ |url-status=live}}</ref> Each subgenus is further divided into sections and subsections.<ref name="Gernandt López García 2005">{{cite journal |last=Gernandt |first=David S. |last2=López |first2=Gretel Geada |last3=García |first3=Sol Ortiz |last4=Liston |first4=Aaron |title=Phylogeny and classification of Pinus |journal=Taxon |volume=54 |issue=1 |date=2005 |doi=10.2307/25065300 |doi-access=free |pages=29–42 |url=https://rc.upr.edu.cu/jspui/bitstream/DICT/2586/1/Geada-LopezTAXON2005.pdf}}</ref>

Many of the smaller groups of ''Pinus'' are composed of closely related species with recent divergence and history of hybridisation. This results in low morphological and genetic differences. This, coupled with low sampling and underdeveloped genetic techniques, has made taxonomy difficult to determine.<ref>{{cite journal |last1=Flores-Rentería |first1=Lluvia |last2=Wegier |first2=Ana |last3=Ortega Del Vecchyo |first3=Diego |last4=Ortíz-Medrano |first4=Alejandra |last5=Piñero |first5=Daniel |last6=Whipple |first6=Amy V. |last7=Molina-Freaner |first7=Francisco |last8=Domínguez |first8=César A. |display-authors=5 |title=Genetic, morphological, geographical and ecological approaches reveal phylogenetic relationships in complex groups, an example of recently diverged pinyon pine species (Subsection Cembroides) |journal=Molecular Phylogenetics and Evolution |volume=69 |issue=3 |pages=940–9 |date=December 2013 |pmid=23831459 |doi=10.1016/j.ympev.2013.06.010 |bibcode=2013MolPE..69..940F }}</ref> Recent research using large genetic datasets has clarified these relationships into the groupings often accepted today.<ref name="Gernandt López García 2005"/>

=== Phylogeny ===

''Pinus'' is the largest genus of the [[Pinaceae]], the pine family, which first appeared in the [[Jurassic]] period.<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=December 2018 |title=Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis |journal=Molecular Phylogenetics and Evolution |volume=129 |pages=106–116 |doi=10.1016/j.ympev.2018.08.011 |pmid=30153503 |bibcode=2018MolPE.129..106R |s2cid=52110440}}</ref> Based on recent [[transcriptome]] analysis, ''Pinus'' is most closely related to the genus ''[[Cathaya]]'', which in turn is closely related to the genus ''[[Picea]]'', the spruces. These genera, with [[fir]]s and [[larch]]es, form the pinoid [[clade]] of the Pinaceae.<ref name=Stull/> Pines first appeared during the Early Cretaceous, with the oldest verified fossil of the genus being ''[[Pinus yorkshirensis]]'' from the [[Hauterivian]]-[[Barremian]] boundary (~130-125 million years ago) from the [[Speeton Clay Formation|Speeton Clay]], England.<ref>{{cite journal |last1=Ryberg |first1=Patricia E. |last2=Rothwell |first2=Gar W. |last3=Stockey |first3=Ruth A. |last4=Hilton |first4=Jason |last5=Mapes |first5=Gene |last6=Riding |first6=James B. |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> However, there are possible records from the Jurassic.<ref>{{cite web |title=Ralikhomo tracksite (zone C/3) (Jurassic of Lesotho) |url=https://paleobiodb.org/classic/basicCollectionSearch?collection_no=149715&is_real_user=1 |website=PBDB.org}}</ref>

The evolutionary history of the genus ''Pinus'' has been complicated by [[Hybrid (biology)|hybridisation]]. Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weak [[reproductive isolation]] make breeding across species more likely.<ref>{{cite journal  |last1=Hernández-León |first1=Sergio |last2=Gernandt |first2=David S. |last3=Pérez de la Rosa |first3=Jorge A. |last4=Jardón-Barbolla |first4=Lev |date=2013-07-30 |title=Phylogenetic relationships and species delimitation in ''Pinus'' section ''Trifoliae'' inferrred from plastid DNA |journal=PLOS ONE |volume=8 |issue=7 |pages=e70501 |bibcode=2013PLoSO...870501H |doi=10.1371/journal.pone.0070501 |pmc=3728320 |pmid=23936218 |doi-access=free}}</ref> As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness.

Two recent phylogenies are given below; the differences between them, and other published phylogenies, demonstrate these complications:
{|class="wikitable"
|-
! colspan=1 |Jin et al. 2021<ref>{{cite journal |title=Phylogenomic and ecological analyses reveal the spatiotemporal evolution of global pines |last1=Jin |first1=Wei-Tao |last2=Gernandt |first2=David S. |last3=Wehenkel |first3=Christian |last4=Xia |first4=Xiao-Mei |last5=Wei |first5=Xiao-Xin |last6=Wang |first6=Xiao-Quan |journal=PNAS |date=May 2021 |doi=10.1073/pnas.2022302118 |volume=118 |issue=20 |pages=e2022302118 |pmid=33941644 |pmc=8157994 |bibcode=2021PNAS..11822302J |doi-access=free}}</ref>
! colspan=1 |Stull et al. 2021<ref name=Stull/>
|- 
|style="vertical-align:top |
{{#invoke:Clade|main |style=font-size:85%;line-height:75%
 |label1=''Pinus''
 |1={{#invoke:Clade|main
  |label1=('''''Strobus''''')
  |1={{#invoke:Clade|main
   |label1=section
   |sublabel1=''Parrya''
   |1={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Nelsonii''
     |1=''[[Pinus nelsonii|P. nelsonii]]''
     |label2=subsection 
     |sublabel2=''Balfouriana''
     |2={{#invoke:Clade|main
      |1=''[[Pinus aristata|P. aristata]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus balfouriana|P. balfouriana]]''
       |2=''[[Pinus longaeva|P. longaeva]]''
       }}
      }}
     }}
    |2={{#invoke:Clade|main
     |label1=subsection 
     |sublabel1=''Cembroides''
     |1={{#invoke:Clade|main
      |1={{#invoke:Clade|main
       |2=''[[Pinus rzedowskii|P. rzedowskii]]''
       |1={{#invoke:Clade|main
        |2=''[[Pinus maximartinezii|P. maximartinezii]]''
        |1=''[[Pinus pinceana|P. pinceana]]''
        }}
       }}
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus quadrifolia|P. quadrifolia]]''
        |2=''[[Pinus monophylla|P. monophylla]]''
        }}
       |2={{#invoke:Clade|main
        |2={{#invoke:Clade|main
         |1=''[[Pinus edulis|P. edulis]]''
         |2=''[[Pinus cembroides|P. cembroides]]''
         }}
        |1={{#invoke:Clade|main
         |2=''[[Pinus remota|P. remota]]''
         |1={{#invoke:Clade|main
          |1=''[[Pinus culminicola|P. culminicola]]''
          |2=''[[Pinus discolor|P. discolor]]''
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   |label2=section 
   |sublabel2=''Quinquefoliae''
   |2={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection 
     |sublabel1=''Gerardiana''
     |1={{#invoke:Clade|main
      |1=''[[Pinus bungeana|P. bungeana]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus squamata|P. squamata]]''
       |2=''[[Pinus gerardiana|P. gerardiana]]''
       }}
      }}
     }}
    |2={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Krempfii''
     |1=''[[Pinus krempfii|P. krempfii]]''
     |label2=subsection
     |sublabel2=''Strobus''
     |2={{#invoke:Clade|main
      |1=''[[Pinus peuce|P. peuce]]''
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus lambertiana|P. lambertiana]]''
        |2={{#invoke:Clade|main
         |1={{#invoke:Clade|main
          |1=''[[Pinus strobus|P. strobus]]''
          |2=''[[Pinus chiapensis|P. chiapensis]]''
          }}
         |2={{#invoke:Clade|main
          |1=''[[Pinus monticola|P. monticola]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus flexilis|P. flexilis]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus strobiformis|P. strobiformis]]''
            |2=''[[Pinus ayacahuite|P. ayacahuite]]''
            }}
           }}
          }}
         }}
        }}
       |2={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus albicaulis|P. albicaulis]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus koraiensis|P. koraiensis]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus sibirica|P. sibirica]]''
           |2=''[[Pinus cembra|P. cembra]]''
           }}
          }}
         }}
        |2={{#invoke:Clade|main
         |1=''[[Pinus parviflora|P. parviflora]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus wallichiana|P. wallichiana]]''
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1={{#invoke:Clade|main
             |2=''[[Pinus dabeshanensis|P. dabeshanensis]]''
             |1=''[[Pinus armandii|P. armandii]]''
             }}
            |2={{#invoke:Clade|main
             |1=''[[Pinus fenzeliana|P. fenzeliana]]''
             |2=''[[Pinus mastersiana|P. mastersiana]]''
             }}
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus bhutanica|P. bhutanica]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus morrisonicola|P. morrisonicola]]''
             |2={{#invoke:Clade|main
              |1=''[[Pinus wangii|P. wangii]]''
              |2={{#invoke:Clade|main
               |1=''[[Pinus kwangtungensis|P. kwangtungensis]]''
               |2=''[[Pinus dalatensis|P. dalatensis]]''
               }}
              }}
             }}
            }}
           }}
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   }}
  |label2=('''''Pinus''''')
  |2={{#invoke:Clade|main
   |label1=section
   |sublabel1=''Pinus''
   |1={{#invoke:Clade|main
    |label1=subsection
    |sublabel1=''Pinaster''
    |1={{#invoke:Clade|main
     |1=''[[Pinus heldreichii|P. heldreichii]]''
     |2={{#invoke:Clade|main
      |1=''[[Pinus pinaster|P. pinaster]]''
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus halepensis|P. halepensis]]''
        |2=''[[Pinus brutia|P. brutia]]''
        }}
       |2={{#invoke:Clade|main
        |1=''[[Pinus pinea|P. pinea]]''
        |2={{#invoke:Clade|main
         |1=''[[Pinus canariensis|P. canariensis]]''
         |2=''[[Pinus roxburghii|P. roxburghii]]''
         }}
        }}
       }}
      }}
     }}
    |label2=subsection
    |sublabel2=''Pinus''
    |2={{#invoke:Clade|main
     |1={{#invoke:Clade|main
      |1=''[[Pinus latteri|P. latteri]]''
      |2=''[[Pinus merkusii|P. merkusii]]''
      }}
     |2={{#invoke:Clade|main
      |1=''[[Pinus resinosa|P. resinosa]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus nigra|P. nigra]]''
       |2={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus densiflora|P. densiflora]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus sylvestris|P. sylvestris]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus mugo|P. mugo]]''
           |2=''[[Pinus uncinata|P. uncinata]]''
           }}
          }}
         }}
        |2={{#invoke:Clade|main
         |1={{#invoke:Clade|main
          |1=''[[Pinus massoniana|P. massoniana]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus thunbergii|P. thunbergii]]''
           |2=''[[Pinus luchuensis|P. luchuensis]]''
           }}
          }}
         |2={{#invoke:Clade|main
          |1={{#invoke:Clade|main
           |1=''[[Pinus kesiya|P. kesiya]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus yunnanensis|P. yunnanensis]]''
            |2=''[[Pinus densata|P. densata]]''
            }}
           }}
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1=''[[Pinus henryi|P. henryi]]''
            |2=''[[Pinus tabuliformis|P. tabuliformis]]''
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus hwangshanensis|P. hwangshanensis]]''
            |2=''[[Pinus taiwanensis|P. taiwanensis]]''
            }}
           }}
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   |label2=section
   |sublabel2=''Trifoliae''
   |2={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Sabiniana''
     |1={{#invoke:Clade|main
      |1=''[[Pinus jeffreyi|P. jeffreyi]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus coulteri|P. coulteri]]''
       |2={{#invoke:Clade|main
        |1=''[[Pinus torreyana|P. torreyana]]''
        |2=''[[Pinus sabiniana|P. sabiniana]]''
        }}
       }}
      }}
     }}
    |2={{#invoke:Clade|main
     |1={{#invoke:Clade|main
      |label1=subsection
      |sublabel1=''Ponderosae''
      |1={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus washoensis|P. washoensis]]''
        |2=''[[Pinus ponderosa|P. ponderosa]]''
        }}
       |2={{#invoke:Clade|main
        |1=''[[Pinus hartwegii|P. hartwegii]]''
        |2={{#invoke:Clade|main
         |1={{#invoke:Clade|main
          |1=''[[Pinus engelmannii|P. engelmannii]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus durangensis|P. durangensis]]''
           |2=''[[Pinus arizonica|P. arizonica]]''
           }}
          }}
         |2={{#invoke:Clade|main
          |1={{#invoke:Clade|main
           |1=''[[Pinus devoniana|P. devoniana]]''
           |2=''[[Pinus montezumae|P. montezumae]]''
           }}
          |2={{#invoke:Clade|main
           |1=''[[Pinus pseudostrobus|P. pseudostrobus]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus gordoniana|P. gordoniana]]''
            |2=''[[Pinus maximinoi|P. maximinoi]]''
            }}
           }}
          }}
         }}
        }}
       }} 
      }}
     |2={{#invoke:Clade|main
      |1={{#invoke:Clade|main
       |label1=subsection
       |sublabel1=''Attenuata''
       |1={{#invoke:Clade|main
        |2=''[[Pinus muricata|P. muricata]]''
        |1={{#invoke:Clade|main
         |1=''[[Pinus radiata|P. radiata]]''
         |2=''[[Pinus attenuata|P. attenuata]]''
         }}
        }}
       }}
      |2={{#invoke:Clade|main
       |label1=subsection
       |sublabel1=''Contorta''
       |1={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus contorta|P. contorta]]''
         |2=''[[Pinus banksiana|P. banksiana]]''
         }}
        |2={{#invoke:Clade|main
         |1=''[[Pinus virginiana|P. virginiana]]''
         |2=''[[Pinus clausa|P. clausa]]''
         }}
        }}
       |label2=subsection
       |sublabel2=''Australes''
       |2={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus pungens|P. pungens]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus serotina|P. serotina]]''
          |2=''[[Pinus rigida|P. rigida]]''
          }}
         }}
        |2={{#invoke:Clade|main
         |1=''[[Pinus glabra|P. glabra]]''
         |2={{#invoke:Clade|main
          |1={{#invoke:Clade|main
           |1=''[[Pinus taeda|P. taeda]]''
           |2=''[[Pinus echinata|P. echinata]]''
           }}
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1=''[[Pinus elliottii|P. elliottii]]''
            |2=''[[Pinus palustris|P. palustris]]''
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus caribaea|P. caribaea]]''
            |2={{#invoke:Clade|main
             |1={{#invoke:Clade|main
              |1=''[[Pinus oocarpa|P. oocarpa]]''
              |2=''[[Pinus tecunumanii|P. tecunumanii]]''
              }}
             |2={{#invoke:Clade|main
              |1={{#invoke:Clade|main
               |1=''[[Pinus greggii|P. greggii]]''
               |2={{#invoke:Clade|main
                |1=''[[Pinus patula|P. patula]]''
                |2=''[[Pinus teocote|P. teocote]]''
                }}
               }}
              |2={{#invoke:Clade|main
               |1={{#invoke:Clade|main
                |1=''[[Pinus lawsonii|P. lawsonii]]''
                |2={{#invoke:Clade|main
                 |1=''[[Pinus herrerae|P. herrerae]]''
                 |2=''[[Pinus pringlei|P. pringlei]]''
                 }}
                }}
               |2={{#invoke:Clade|main
                |1={{#invoke:Clade|main
                 |1=''[[Pinus leiophylla|P. leiophylla]]''
                 |2=''[[Pinus lumholtzii|P. lumholtzii]]''
                 }}
                |2={{#invoke:Clade|main
                 |1={{#invoke:Clade|main
                  |1=''[[Pinus praetermissa|P. praetermissa]]''
                  |2=''[[Pinus jaliscana|P. jaliscana]]''
                  }}
                 |2={{#invoke:Clade|main
                  |1=''[[Pinus luzmariae|P. luzmariae]]''
                  |2=''[[Pinus georginae|P. georginae]]''
                  }}
                 }}
                }}
               }}
              }}
             }}
            }}
           }}
          }} 
         }}
        }}
       }}
      }}
     }}
    }}
   }}
  }}
}}
|
{{#invoke:Clade|main |style=font-size:85%;line-height:75%
 |label1='''''Pinus'''''
 |1={{#invoke:Clade|main
  |label1=('''''Strobus''''')
  |1={{#invoke:Clade|main
   |label1=section
   |sublabel1='''''Parrya'''''
   |1={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Nelsonii''
     |1=''[[Pinus nelsonii|P. nelsonii]]''
     |label2=subsection
     |sublabel2=''Balfouriana''
     |2={{#invoke:Clade|main
      |1=''[[Pinus aristata|P. aristata]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus balfouriana|P. balfouriana]]''
       |2=''[[Pinus longaeva|P. longaeva]]''
       }}
      }}
     }}
    |2={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Cembroides''
     |1={{#invoke:Clade|main
      |1={{#invoke:Clade|main
       |1=''[[Pinus monophylla|P. monophylla]]''
       |2={{#invoke:Clade|main
        |1=''[[Pinus juarezensis|P. juarezensis]]''
        |2=''[[Pinus quadrifolia|P. quadrifolia]]''
        }}
       }}
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |2=''[[Pinus maximartinezii|P. maximartinezii]]''
        |1=''[[Pinus pinceana|P. pinceana]]''
        }}
       |2={{#invoke:Clade|main
        |1=''[[Pinus rzedowskii|P. rzedowskii]]''
        |2={{#invoke:Clade|main
         |1=''[[Pinus edulis|P. edulis]]''
         |2={{#invoke:Clade|main
          |1={{#invoke:Clade|main
           |1=''[[Pinus discolor|P. discolor]]''
           |2=''[[Pinus johannis|P. johannis]]''
           }}
          |2={{#invoke:Clade|main
           |1=''[[Pinus culminicola|P. culminicola]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus cembroides|P. cembroides]]''
            |2=''[[Pinus remota|P. remota]]''
            }}
           }}
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   |label2=section
   |sublabel2='''''Quinquefoliae'''''
   |2={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Gerardiana''
     |1={{#invoke:Clade|main
      |1=''[[Pinus bungeana|P. bungeana]]''
      |2=''[[Pinus gerardiana|P. gerardiana]]''
      }}
     }}
    |2={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Krempfii''
     |1=''[[Pinus krempfii|P. krempfii]]''
     |label2=subsection
     |sublabel2=''Strobus''
     |2={{#invoke:Clade|main
      |1={{#invoke:Clade|main
       |label1=series
       |sublabel1=''Macedonia''
       |1=''[[Pinus peuce|P. peuce]]''
       }}
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |label1=series
        |sublabel1=''Strobi''
        |1={{#invoke:Clade|main
         |1=''[[Pinus strobus|P. strobus]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus flexilis|P. flexilis]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus ayacahuite|P. ayacahuite]]''
           |2=''[[Pinus strobiformis|P. strobiformis]]''
          }}
          }}
         }}
        }}
       |2={{#invoke:Clade|main
        |label1=series
        |sublabel1=''Cembrae''
        |1={{#invoke:Clade|main
         |1=''[[Pinus parviflora|P. parviflora]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus monticola|P. monticola]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus cembra|P. cembra]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus lambertiana|P. lambertiana]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus albicaulis|P. albicaulis]]''
             |2=''[[Pinus dalatensis|P. dalatensis]]''
             }}
            }}
           }}
          }}
         }}
        |label2=series
        |sublabel2=''Pumilae''
        |2={{#invoke:Clade|main
         |1=''[[Pinus pumila|P. pumila]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus wallichiana|P. wallichiana]]''
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1=''[[Pinus koraiensis|P. koraiensis]]''
            |2=''[[Pinus sibirica|P. sibirica]]''
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus squamata|P. squamata]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus morrisonicola|P. morrisonicola]]''
             |2={{#invoke:Clade|main
              |1=''[[Pinus armandii|P. armandii]]''
              |2={{#invoke:Clade|main
               |1=''[[Pinus fenzeliana|P. fenzeliana]]''
               |2={{#invoke:Clade|main
                |1=''[[Pinus kwangtungensis|P. kwangtungensis]]''
                |2={{#invoke:Clade|main
                 |1=''[[Pinus bhutanica|P. bhutanica]]''
                 |2=''[[Pinus wangii|P. wangii]]''
                 }}
                }}
               }}
              }}
             }}
            }}
           }}
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   }}
  |label2=('''''Pinus''''')
  |2={{#invoke:Clade|main
   |label1=section
   |sublabel1=''Pinus''
   |1={{#invoke:Clade|main
    |label1=subsection
    |sublabel1=''Pinaster''
    |1={{#invoke:Clade|main
     |1=''[[Pinus heldreichii|P. heldreichii]]''
     |2={{#invoke:Clade|main
      |1={{#invoke:Clade|main
       |1=''[[Pinus halepensis|P. halepensis]]''
       |2=''[[Pinus brutia|P. brutia]]''
       }}
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus pinaster|P. pinaster]]''
        |2=''[[Pinus pinea|P. pinea]]''
        }}
       |2={{#invoke:Clade|main
        |1=''[[Pinus canariensis|P. canariensis]]''
        |2=''[[Pinus roxburghii|P. roxburghii]]''
        }}
       }}
      }}
     }}
    |label2=subsection
    |sublabel2=''Pinus''
    |2={{#invoke:Clade|main
     |1={{#invoke:Clade|main
      |label1=series
      |sublabel1=''Pinus''
      |1={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |1=''[[Pinus densata|P. densata]]''
        |2={{#invoke:Clade|main
         |1=''[[Pinus latteri|P. latteri]]''
         |2=''[[Pinus kesiya|P. kesiya]]''
         }}
        }}
       |2={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus nigra|P. nigra]]''
         |2=''[[Pinus resinosa|P. resinosa]]''
         }}
        |2={{#invoke:Clade|main
         |1=''[[Pinus sylvestris|P. sylvestris]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus mugo|P. mugo]]''
          |2=''[[Pinus uncinata|P. uncinata]]''
          }}
         }}
        }}
       }}
      }}
     |2={{#invoke:Clade|main
      |label1=series
      |sublabel1=''Tropicales''
      |1={{#invoke:Clade|main
       |1=''[[Pinus tropicalis|P. tropicalis]]''
       |2={{#invoke:Clade|main
        |1=''[[Pinus hwangshanensis|P. hwangshanensis]]''
        |2={{#invoke:Clade|main
         |1=''[[Pinus massoniana|P. massoniana]]''
         |2=''[[Pinus merkusii|P. merkusii]]''
         }}
        }}
       }}
      |label2=series
      |sublabel2=''Tabuliformes''
      |2={{#invoke:Clade|main
       |1=''[[Pinus densiflora|P. densiflora]]''
       |2={{#invoke:Clade|main
        |1={{#invoke:Clade|main
         |1=''[[Pinus thunbergii|P. thunbergii]]''
         |2=''[[Pinus taiwanensis|P. taiwanensis]]''
         }}
        |2={{#invoke:Clade|main
         |1=''[[Pinus yunnanensis|P. yunnanensis]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus luchuensis|P. luchuensis]]''
          |2=''[[Pinus tabuliformis|P. tabuliformis]]''
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   |label2=section
   |sublabel2=''Trifoliae''
   |2={{#invoke:Clade|main
    |1={{#invoke:Clade|main
     |label1=subsection
     |sublabel1=''Contorta''
     |1={{#invoke:Clade|main
      |1=''[[Pinus virginiana|P. virginiana]]''
      |2={{#invoke:Clade|main
       |1=''[[Pinus contorta|P. contorta]]''
       |2=''[[Pinus clausa|P. clausa]]''
       }}
      }}
     }}
    |2={{#invoke:Clade|main
     |1={{#invoke:Clade|main
      |label1=subsection
      |sublabel1=''Ponderosae''
      |1={{#invoke:Clade|main
       |1=''[[Pinus maximinoi|P. maximinoi]]''
       |2={{#invoke:Clade|main
        |1=''[[Pinus montezumae|P. montezumae]]''
        |2={{#invoke:Clade|main
         |1={{#invoke:Clade|main
          |1=''[[Pinus sabiniana|P. sabiniana]]''
          |2=''[[Pinus torreyana|P. torreyana]]''
          }}
         |2={{#invoke:Clade|main
          |1=''[[Pinus ponderosa|P. ponderosa]]''
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1=''[[Pinus gordoniana|P. gordoniana]]''
            |2=''[[Pinus yecorensis|P. yecorensis]]''
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus cooperi|P. cooperi]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus durangensis|P. durangensis]]''
             |2={{#invoke:Clade|main
              |1=''[[Pinus nubicola|P. nubicola]]''
              |2={{#invoke:Clade|main
               |1=''[[Pinus coulteri|P. coulteri]]''
               |2={{#invoke:Clade|main
                |1={{#invoke:Clade|main
                 |1=''[[Pinus jeffreyi|P. jeffreyi]]''
                 |2=''[[Pinus washoensis|P. washoensis]]''
                 }}
                |2={{#invoke:Clade|main
                 |1=''[[Pinus arizonica|P. arizonica]]''
                 |2={{#invoke:Clade|main
                  |1=''[[Pinus engelmannii|P. engelmannii]]''
                  |2={{#invoke:Clade|main
                   |1=''[[Pinus devoniana|P. devoniana]]''
                   |2={{#invoke:Clade|main
                    |1=''[[Pinus hartwegii|P. hartwegii]]''
                    |2=''[[Pinus pseudostrobus|P. pseudostrobus]]''
                    }}
                   }}
                  }}
                 }}
                }}
               }}
              }}
             }}
            }}
           }}
          }}
         }}
        }}
       }} 
      }}
     |2={{#invoke:Clade|main
      |label1=subsection
      |sublabel1=''Attenuata''
      |1={{#invoke:Clade|main
       |label1=series
       |sublabel1=''Leiophyllae''
       |1={{#invoke:Clade|main
        |1=''[[Pinus patula|P. patula]]''
        |2=''[[Pinus leiophylla|P. leiophylla]]''
        }}
       |label2=series
       |sublabel2=''Attenuatae''
       |2={{#invoke:Clade|main
        |1=''[[Pinus tecunumanii|P. tecunumanii]]''
        |2={{#invoke:Clade|main
         |1=''[[Pinus muricata|P. muricata]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus radiata|P. radiata]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus attenuata|P. attenuata]]''
           |2=''[[Pinus glabra|P. glabra]]''
           }}
          }}
         }}
        }}
       }}
      |label2=subsection
      |sublabel2=''Australes''
      |2={{#invoke:Clade|main
       |1={{#invoke:Clade|main
        |label1=series
        |sublabel1=''Palustres''
        |1={{#invoke:Clade|main
         |1=''[[Pinus herrerae|P. herrerae]]''
         |2={{#invoke:Clade|main
          |1=''[[Pinus caribaea|P. caribaea]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus palustris|P. palustris]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus echinata|P. echinata]]''
            |2=''[[Pinus occidentalis|P. occidentalis]]''
            }}
           }}
          }}
         }}
        }}
       |2={{#invoke:Clade|main
        |1=''[[Pinus praetermissa|P. praetermissa]]''
        |2={{#invoke:Clade|main
         |label1=series
         |sublabel1=''Taeda''
         |1={{#invoke:Clade|main
          |1=''[[Pinus serotina|P. serotina]]''
          |2={{#invoke:Clade|main
           |1=''[[Pinus rigida|P. rigida]]''
           |2={{#invoke:Clade|main
            |1=''[[Pinus pungens|P. pungens]]''
            |2=''[[Pinus taeda|P. taeda]]''
            }}
           }}
          }}
         |label2=series
         |sublabel2=''Oocarpae''
         |2={{#invoke:Clade|main
          |1=''[[Pinus elliottii|P. elliottii]]''
          |2={{#invoke:Clade|main
           |1={{#invoke:Clade|main
            |1=''[[Pinus teocote|P. teocote]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus greggii|P. greggii]]''
             |2=''[[Pinus lumholtzii|P. lumholtzii]]''
             }}
            }}
           |2={{#invoke:Clade|main
            |1=''[[Pinus jaliscana|P. jaliscana]]''
            |2={{#invoke:Clade|main
             |1=''[[Pinus luzmariae|P. luzmariae]]''
             |2={{#invoke:Clade|main
              |1=''[[Pinus georginae|P. georginae]]''
              |2={{#invoke:Clade|main
               |1=''[[Pinus maestrensis|P. maestrensis]]''
               |2={{#invoke:Clade|main
                |1=''[[Pinus pringlei|P. pringlei]]''
                |2={{#invoke:Clade|main
                 |1=''[[Pinus lawsonii|P. lawsonii]]''
                 |2={{#invoke:Clade|main
                  |1=''[[Pinus cubensis|P. cubensis]]''
                  |2=''[[Pinus oocarpa|P. oocarpa]]''
                  }}
                 }}
                }}
               }}
              }}
             }}
            }}
           }}
          }} 
         }}
        }}
       }}
      }}
     }}
    }}
   }}
  }}
}}
|}