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FOC | Family List | FOC Vol. 4 | Pinaceae

1. Pinus Linnaeus, Sp. Pl. 2: 1000. 1753.

松属 song shu

Trees or rarely shrubs, evergreen, with regularly whorled branches; branchlets strongly dimorphic: long branchlets bearing scalelike leaves and spreading leaf bundles; short branchlets bearing leaves in bundles of 2-5(-7); winter buds large, with numerous scales. Leaves needlelike, slender or stout, straight or twisted, triangular, flabellate-triangular, or semiorbicular in cross section, stomatal lines several, on 1, 2, or all surfaces, vascular bundles 1 or 2, resin canals 2-10 or more, marginal or median, rarely internal, base enclosed by persistent or deciduous, membranous sheath. Pollen cones usually borne in spikelike clusters at base of 1st-year branchlets, sessile, cylindric or ovoid; pollen 2-saccate. Seed cones pedunculate or subsessile, erect or pendulous, cylindric or ovoid, maturing in 2nd or 3rd year. Seed scales spirally arranged, woody, exposed apex thickened and ridged (the apophysis), with a prominent protuberance (umbo), usually terminating in a spine or prickle, persistent. Bracts minute. Seeds variable in color, shape, and size, winged or not; wing adnate or articulated to seed. Cotyledons 3-18. Germination epigeal. 2n = 24*.

About 110 species: N Africa, Asia, Europe, North America; 39 species (seven endemic, 16 introduced) in China.


1 Needles with basal sheath deciduous and basal scalelike leaves not decurrent, cross section with 1 vascular bundle; umbo terminal or dorsal.   (2)
+ Needles with basal sheath persistent and basal scalelike leaves decurrent, cross section with 2 vascular bundles; umbo dorsal.   (16)
       
2 (1) Umbo dorsal; needles 3-5 per bundle.   (3)
+ Umbo terminal; needles (2-)5 per bundle.   (5)
       
3 (2) Needles 4 or 5 per bundle; bark dark brown, flaking   25 Pinus squamata
+ Needles 3 per bundle; bark white, irregularly flaking.   (4)
       
4 (3) Seed cones 12-20 cm; seeds cylindric, ca. 2.5 cm   26 Pinus gerardiana
+ Seed cones 5-7 cm; seeds ovoid, ca. 1 cm   27 Pinus bungeana
       
5 (2) Seeds wingless or only shortly winged.   (6)
+ Seeds long winged.   (10)
       
6 (5) Seed cones dehiscent at maturity; branchlets glabrous.   (7)
+ Seed cones indehiscent or imperfectly dehiscent; branchlets densely pubescent.   (8)
       
7 (6) Seeds wingless or ridged along abaxial margin; seed coat thick; apophyses not or slightly reflexed.at margin   31 Pinus armandi
+ Seeds with a rudimentary wing 2-7 mm; seed coat thin; apophyses obviously reflexed at margin   32 Pinus fenzeliana
       
8 (6) Needles slender, 4-8 cm × 0.5-1 mm; resin canals 2, marginal; shrubs   28 Pinus pumila
+ Needles stout, 7-12 cm × 1-1.5 mm; resin canals 3, median; trees.   (9)
       
9 (8) First-year branchlets with red-brown hairs; needles with 6-8 blue-gray stomatal lines along each abaxial surface; seed cones 9-14 cm; seed scales acuminate at apex, recurved   29 Pinus koraiensis
+ First-year branchlets with yellow-brown hairs; needles with 3-5 gray-white stomatal lines along each abaxial surface; seed cones 5-8 cm; seed scales rounded at apex, incurved   30 Pinus sibirica
       
10 (5) Needles 7-20 cm; seed cones cylindric to narrowly cylindric, 7-25 cm.   (11)
+ Needles less than 8 cm; seed cones ovoid, ovoid-ellipsoid, or ellipsoid-cylindric, usually less than 10 cm.   (13)
       
11 (10) Needles not pendulous, 6-14 cm; seed cones 7-20 cm   38 Pinus strobus
+ Needles pendulous, (6-)11-24 cm; seed cones 10-30 cm.   (12)
       
12 (11) Needles dark green adaxially; 1st-year branchlets green with faint bloom, glabrous; abaxial resin canal of needles always median   34 Pinus wallichiana
+ Needles grass green adaxially, conspicuously whitish bloomed on 2 abaxial surfaces; 1st-year branchlets conspicuously whitish bloomed, glandular pubescent; 1(or 2) abaxial resin canal(s) of needles always asymmetrically placed   33 Pinus bhutanica
       
13 (10) Needles less than 1 mm in diam.; seed cones sessile or shortly pedunculate.   (14)
+ Needles 1-1.5 mm in diam.; seed cones distinctly pedunculate.   (15)
       
14 (13) Branchlets densely pubescent; seed cones sessile; seeds as long as broad wing; needles 3.5-5.5 cm   39 Pinus parviflora
+ Branchlets glabrous or puberulent; seed cones shortly pedunculate; seeds ca. 1/2 as long as narrow wing; needles 4-8 cm   35 Pinus morrisonicola
       
15 (13) Branchlets densely pubescent; resin canals 3, median   36 Pinus wangii
+ Branchlets glabrous, rarely puberulent; resin canals 2, marginal, sometimes also 1 median   37 Pinus kwangtungensis
       
16 (1) Wing adnate to seed; needles 3 per bundle; apophyses distinctly protruded, with transverse ridges and spiny umbo   1 Pinus roxburghii
+ Wing articulated to seed; needles 2-5 per bundle; apophyses and umbo of different types.   (17)
       
17 (16) Needles (2 or)3(-5) per bundle.   (18)
+ Needles 2(or 3) per bundle.   (28)
       
18 (17) Trunk and branches usually with adventitious sprouts; needles mostly to 20 cm.   (19)
+ Trunk and branches usually without adventitious sprouts; needles mostly 15-45 cm.   (21)
       
19 (18) Needles 2(or 3) per bundle, ca. 1 mm in diam.; winter buds 0.5-0.7(-1) cm in diam.; bark plates with evident resin pockets; adaxial surface of seed scales lacking contrasting border distally   11 Pinus echinata
+ Needles 3(-5) per bundle, 1-1.5(-2) mm in diam.; winter buds 1-1.5(-2) cm in diam.; bark plates without evident resin pockets; adaxial surface of seed scales with dark red-brown border distally.   (20)
       
20 (19) Needles (12-)15-21 cm, slightly twisted; branchlets initially orange or yellow-orange, often glaucous; seed cones late dehiscent after maturity   12 Pinus serotina
+ Needles 5-10(-15) cm, twisted; branchlets initially orange-brown, not glaucous; seed cones dehiscent at maturity   13 Pinus rigida
       
21 (18) Needles slender, 10-30 cm × 0.7-1.2 mm (except P. yunnanensis var. pygmaea with stout needles 5-13 cm).   (22)
+ Needles ± stout, 12-45 cm × 1.5-2 mm.   (24)
       
22 (21) Umbo blunt or shortly mucronate in apical part of cone; needles 2(or 3) per bundle, 12-20 cm, slightly twisted   2 Pinus massoniana
+ Umbo shortly mucronate; needles (2 or)3 per bundle, 10-30 cm, not twisted.   (23)
       
23 (22) Needles slender and pliant, less than 1 mm in diam.; resin canals marginal; apophyses ± pyramidal, distinctly cross keeled   3 Pinus kesiya
+ Needles relatively stout and stiff (slender and pliant in plants growing in dry, hot valleys), 1-1.2 mm in diam.; resin canals marginal and median; apophyses swollen, usually not pyramidal, not or slightly cross keeled   4 Pinus yunnanensis
       
24 (21) Winter buds 3-4 cm, silvery white; seed cones 15-25 cm   14 Pinus palustris
+ Winter buds to 2 cm, reddish, chestnut, orange, or silvery brown; seed cones usually 5-15 cm.   (25)
       
25 (24) Seed cones pedunculate.   (26)
+ Seed cones sessile or subsessile.   (27)
       
26 (25) Seed cones (7-)9-18(-20) cm, all deciduous; umbo central, depressed-pyramidal, with a stout, short prickle; needles 2 or 3 per bundle   15 Pinus elliottii
+ Seed cones 5-10(-12) cm, often a few basal cones persistent; umbo slightly projecting, ending in a straight, minute prickle; needles (2 or)3(-5) per bundle   16 Pinus caribaea
       
27 (25) Branchlets ca. 1 cm in diam.; terminal bud lanceolate-cylindric, mostly less than 1 cm wide, slightly resinous; seed cones mostly dull yellow-brown, narrowly ovoid when open   17 Pinus taeda
+ Branchlets 1-2 cm in diam.; terminal bud ovoid, ca. 1 cm wide, very resinous; seed cones mostly reddish brown, broadly ovoid when open   18 Pinus ponderosa
       
28 (17) Needles mostly 20-30 cm or more.   (29)
+ Needles mostly less than 20 cm.   (33)
       
29 (28) Umbo blunt or with sunken, short mucro.   (30)
+ Umbo ± spiny or pointed at apex.   (31)
       
30 (29) Needle with 2 resin canals; seed cones red-brown at maturity   6 Pinus latteri
+ Needle with up to 9 resin canals; seed cones pale brown at maturity   19 Pinus tropicalis
       
31 (29) Seed cones borne on stalks to 3 cm, chocolate brown; apophyses lustrous   15 Pinus elliottii
+ Seed cones sessile or subsessile, brown, yellow-, or red-brown; apophyses rarely lustrous.   (32)
       
32 (31) Branchlets ca. 1 cm in diam.; terminal bud lanceolate-cylindric, usually less than 1 cm wide, slightly resinous; seed cones usually dull yellow-brown, narrowly ovoid when open   17 Pinus taeda
+ Branchlets 1-2 cm in diam.; terminal bud ovoid, ca. 1 cm wide, very resinous; seed cones usually reddish brown, broadly ovoid when open   18 Pinus ponderosa
       
33 (28) Seed cones 9-18 cm   20 Pinus pinaster
+ Seed cones usually not more than 8 cm.   (34)
       
34 (33) Needles slender and pliant, ca. 1 mm in diam.; apophyses relatively thin, flat or slightly raised.   (35)
+ Needles stout, 1-2 mm in diam.; apophyses ± swollen or pyramidal.   (37)
       
35 (34) Trunk and branches usually with adventitious sprouts; umbo with a stout, sharp prickle   11 Pinus echinata
+ Trunk and branches usually without adventitious sprouts; umbo flat or blunt.   (36)
       
36 (35) Needles 12-20 cm, slightly twisted   2 Pinus massoniana
+ Needles 5-12 cm, straight 7. P. densiflora   (34)
       
37 (34) Needles shorter, longest ones not more than 8 cm, strongly twisted.   (38)
+ Needles (5-)9-16 cm, straight or rarely slightly twisted.   (40)
       
38 (37) Seed cones strongly asymmetric, curved forward on branches; needles 2-5 cm   23 Pinus banksiana
+ Seed cones symmetric or nearly so; needles usually longer.   (39)
       
39 (38) Bark on distal sections of trunk orange, plated; needles blue-, gray-, or yellow-green, stomatal lines conspicuous; branchlets initially dull green or orange-brown, not glaucous; adaxial surface of seed scales lacking contrasting border distally   10 Pinus sylvestris
+ Bark on distal sections of trunk reddish, scaly; needles deep to pale yellow-green, stomatal lines inconspicuous; branchlets initially red or purple tinged, often glaucous; adaxial surface of seed scales with strong purple-red or -brown border distally   24 Pinus virginiana
       
40 (37) Seed cones deciduous.   (41)
+ Seed cones persistent.   (42)
       
41 (40) Winter buds silvery white, cylindric-ellipsoid or cylindric   21 Pinus thunbergii
+ Winter buds brown, ovoid or oblong-ovoid   22 Pinus nigra
       
42 (40) Needle with resin canals median, rarely also marginal; needles 4.5-17 cm; seed cones 3-6 cm   8 Pinus taiwanensis
+ Needle with resin canals marginal, occasionally 1 or 2 median; needles 6-15 cm; seed cones 2.5-9 cm.   (43)
       
43 (42) Bark initially reddish brown, becoming dark grayish brown; 1st-year branchlets shiny yellowish brown; needle sheaths initially 0.5-1 cm; resin canals 3 or 4; pollen cones 1-1.8 cm; seed cones shiny chocolate brown at maturity   5 Pinus densata
+ Bark dull grayish brown or dark gray; 1st-year branchlets light grayish brown, light brown, or yellowish brown; needle sheaths initially 1-2 cm; resin canals 5-9; pollen cones 0.5-0.9 cm; seed cones yellowish brown to brown at maturity   9 Pinus tabuliformis

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