647: in) 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 milligrams are seemingly adapted for wind dispersal. Pines having seeds larger than 100 mg are more likely to have benefited from adaptations that promote animal dispersal, particularly by birds. Pinaceae that persist in areas where
1894:
1906:
131:
102:
554:
1842:). They are produced and stored in specialized secretory areas known as resin ducts, resin blisters, or resin cavities. Resins have the ability to wash away, trap, fend off antagonists, and are also involved in wound sealing. They are an effective defense mechanism because they have toxic and inhibitory effects on invaders, such as insects or pathogens. Resins could have developed as an evolutionary defense against
664:
680:
699:
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,
695:
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
735:): 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.
3433:
Fäldt, Jenny; Martin, Diane; Miller, Barbara; Rawat, Suman; Bohlmann, Jörg (2003-01-01). "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".
654:
Boreal conifers have many adaptions for winter. The narrow conical shape of northern conifers, and their downward-drooping limbs help them shed snow, and many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening".
760:): 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.
803:): 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.
1783:
and resins. Constitutive defenses are always expressed and offer immediate protection from invaders but could also be defeated by antagonists that have evolved adaptations to these defense mechanisms. One of the common secondary compounds used by
1886:
species. It has been found that MJ stimulated the activation of PP cells and formation of xylem traumatic resin ducts (TD). These are structures that are involved in the release of phenolics and resins, both forms of defense mechanism.
2264:
Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu; Smith, Stephen A.; Yi, Ting-Shuang; et al. (2021).
3314:
Nagy, Nina E.; Franceschi, Vincent R.; Solheim, Halvor; Krekling, Trygve; Christiansen, Erik (2000-03-01). "Wound-induced traumatic resin duct development in stems of Norway spruce (Pinaceae): anatomy and cytochemical traits".
696:
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.
3062:
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
1710:
appears to be non-monophyletic, containing many disparately related members of
Pinaceae. While Pinaceae, and indeed all of its subfamilies, substantially predate the break up of the super-continent
2798:
Patricia E. Ryberg; Gar W. Rothwell; Ruth A. Stockey; Jason Hilton; Gene Mapes; James B. Riding (2012). "Reconsidering
Relationships among Stem and Crown Group Pinaceae: Oldest Record of the Genus
2374:
Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (July 19, 2021).
722:
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.
2331:
1930:
Gernandt, David S.; Holman, Garth; Campbell, Christopher; Parks, Matthew; Mathews, Sarah; Raubeson, Linda A.; Liston, Aaron; Stockey, Ruth A.; Rothwell, Gar W. (September 2016).
1718:. 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.
714:): 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
2635:
Rothwell, Gar W.; Mapes, Gene; Stockey, Ruth A.; Hilton, Jason (April 2012). "The seed cone
Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae".
2223:
3557:
1818:
is resins. Resins are also one of the primary defenses used against attack. Resins are short term defenses that are composed of a complex combination of volatile
1776:
1761:
1779:
are typically the first line of defenses used against antagonists and can include sclerified cells, lignified periderm cells, and secondary compounds such as
2728:
Blokhina, N. I.; Afonin, M. (2007). "Fossil wood Cedrus penzhinaensis sp. nov. (Pinaceae) from the Lower
Cretaceous of north-western Kamchatka (Russia)".
2083:
Robert A. Price, Jeanine Olsen-Stojkovich & Jerold M. Lowenstein (1987). "Relationships among the genera of
Pinaceae: an immunological comparison".
1650:
3144:"Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection"
2439:"Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms"
1694:
2183:
Leslie, Andrew B.; Beaulieu, Jeremy; Holman, Garth; Campbell, Christopher S.; Mei, Wenbin; Raubeson, Linda R.; Mathews, Sarah; et al. (2018).
515:, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of
4886:
2675:"A new species of Pityostrobus (Pinaceae) from the Cretaceous of California: moving towards understanding the Cretaceous radiation of Pinaceae"
1706:
1700:
1760:
defenses are prevalent in the bark of the trees. This part of the tree contributes a complex defensive boundary against external antagonists.
1676:, 157.3-154.7 million years ago) of Scotland, which likely belongs to the pinoid grouping of the family. Pinaceae rapidly radiated during the
4925:
3550:
1893:
1746:
have evolved a myriad of mechanical and chemical defenses, or a combination of the two, in order to protect themselves against antagonists.
503:
trait and genetic analysis. They are the largest extant conifer family in species diversity, with between 220 and 250 species (depending on
1742:
attack which often leads to tree death. In order to combat these stresses, trees need to adapt or evolve defenses against these stresses.
1808:
1765:
2489:
Leslie, Andrew B.; Beaulieu, Jeremy; Holman, Garth; Campbell, Christopher S.; Mei, Wenbin; Raubeson, Linda R.; Mathews, Sarah (2018).
2546:"New species of the genus Schizolepidopsis (conifers) from the Albian of the Russian high Arctic and geological history of the genus"
608:, with subopposite or whorled branches, and spirally arranged, linear (needle-like) leaves. The embryos of Pinaceae have three to 24
4834:
3605:
3543:
4899:
4847:
4986:
3515:
3088:"Induction of anatomically based defense responses in stems of diverse conifers by methyl jasmonate: a phylogenetic perspective"
2594:
Matsunaga, Kelly K. S.; Herendeen, Patrick S.; Herrera, Fabiany; Ichinnorov, Niiden; Crane, Peter R.; Shi, Gongle (2021-05-10).
2837:
Leslie, Andrew B.; Beaulieu, Jeremy M.; Rai, Hardeep S.; Crane, Peter R.; Donoghue, Michael J.; Mathews, Sarah (2012-10-02).
1985:
4904:
619:
are large and usually woody, 2–60 centimetres (1–24 inches) long, with numerous spirally arranged scales, and two winged
3519:
3375:"Defense Mechanisms of Conifers: Differences in Constitutive and Wound-Induced Monoterpene Biosynthesis Among Species"
1905:
5048:
1616:
Multiple molecular studies indicate that in contrast to previous classifications placing it outside the conifers,
4930:
4873:
1882:(MJ) as an antagonist. Methyl jasmonate is known to be able to induce defense responses in the stems of multiple
5012:
2904:
Cherubini, Paolo; Fontana, Giovanni; Rigling, Daniel; Dobbertin, Matthias; Brang, Peter; Innes, John L. (2002).
5043:
4912:
3584:
1654:
likely represent stem-group members of the
Pinaceae, the first good records of which are in the Middle-Late
3610:
3600:
3535:
2376:"Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms"
2267:"Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms"
4730:
1620:
may in fact be the sister group to the
Pinaceae, with both lineages having diverged during the early-mid
2032:
5007:
4826:
4800:
3525:
2761:
1977:
4999:
5089:
4940:
4852:
3505:
2757:"A Lower Cretaceous (Valanginian) seed cone provides the earliest fossil record for Picea (Pinaceae)"
2674:
2121:
700:
based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves:
130:
2490:
2360:
2082:
2033:"Wind dispersal capacity of pine seeds and the evolution of different seed dispersal modes in pines"
3595:
3260:"Anatomical-based defense responses of Scots pine (Pinus sylvestris) stems to two fungal pathogens"
2596:"Ovulate Cones of Schizolepidopsis ediae sp. nov. Provide Insights into the Evolution of Pinaceae"
1811:
responses need to be activated by certain cues, such as herbivore damage or other biotic signals.
4821:
2906:"Tree-Life History Prior to Death: Two Fungal Root Pathogens Affect Tree-Ring Growth Differently"
2545:
1878:
is a very active area of study with numerous studies being conducted. Many of these studies use
4966:
4795:
4683:
1863:
2960:"Terpenoid biosynthesis and specialized vascular cells of conifer defense. - Semantic Scholar"
1666:(descendant of the last common ancestor of all living species) member of Pinaceae is the cone
4961:
4891:
2347:
4953:
2797:
1931:
455:
trees or shrubs, including many of the well-known conifers of commercial importance such as
4756:
3511:
2917:
2850:
2686:
2557:
2387:
2286:
2278:
2133:
2049:
1932:"Phylogenetics of extant and fossil Pinaceae: methods for increasing topological stability"
500:
3007:
Franceschi, Vincent R.; Krokene, Paal; Christiansen, Erik; Krekling, Trygve (2005-08-01).
1866:
in tree stems, roots, and leaves. Oleoresin is also needed in order to classify conifers.
8:
668:
534:
512:
4813:
4787:
2921:
2854:
2690:
2561:
2391:
2282:
2137:
2053:
1726:
External stresses on plants have the ability to change the structure and composition of
5056:
4808:
4721:
3590:
3467:
3407:
3374:
3340:
3009:"Anatomical and chemical defenses of conifer bark against bark beetles and other pests"
2989:
2935:
2881:
2838:
2819:
2737:
2710:
2573:
2526:
2463:
2438:
2419:
2312:
2165:
2102:
2065:
504:
346:
125:
2673:
Smith, Selena Y.; Stockey, Ruth A.; Rothwell, Gar W.; Little, Stefan A. (2017-01-02).
5084:
4948:
4743:
4366:
3459:
3451:
3412:
3394:
3348:
3332:
3289:
3281:
3237:
3229:
3173:
3165:
3117:
3109:
3038:
3030:
3025:
3008:
2981:
2976:
2959:
2930:
2905:
2886:
2868:
2780:
2702:
2652:
2617:
2577:
2518:
2510:
2468:
2423:
2411:
2403:
2375:
2316:
2304:
2266:
2246:
2204:
2157:
2149:
2085:
1981:
1951:
3530:
3471:
2993:
2823:
2741:
2714:
2530:
2339:
2169:
5061:
4748:
4406:
3567:
3443:
3402:
3386:
3324:
3271:
3219:
3209:
3155:
3099:
3020:
2971:
2925:
2876:
2858:
2811:
2770:
2694:
2644:
2607:
2565:
2502:
2491:"An overview of extant conifer evolution from the perspective of the fossil record"
2458:
2450:
2395:
2335:
2296:
2238:
2196:
2185:"An overview of extant conifer evolution from the perspective of the fossil record"
2141:
2122:"Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis"
2094:
2057:
1943:
1879:
1751:
1677:
397:
2698:
2569:
4976:
4735:
4552:
4449:
4260:
3880:
3849:
2120:
Ran, Jin-Hua; Shen, Ting-Ting; Wu, Hui; Gong, Xun; Wang, Xiao-Quan (2018-12-01).
2040:
1632:
Pinaceae is estimated to have diverged from other conifer groups during the late
538:
4917:
3276:
3259:
3104:
3087:
2145:
2002:
1692:(cedar) first appear during the Early Cretaceous. The extinct Cretaceous genera
5035:
4642:
4517:
4442:
3160:
3143:
2399:
2300:
1727:
525:
520:
155:
3447:
3214:
3197:
2291:
5078:
4706:
4635:
4593:
4570:
4545:
4494:
4476:
4386:
4197:
4137:
4123:
4072:
4038:
3764:
3455:
3398:
3336:
3285:
3233:
3198:"Physiological trade-offs in the complexity of pine tree defensive chemistry"
3169:
3113:
3034:
2872:
2706:
2621:
2514:
2407:
2153:
1955:
1859:
1827:
1750:
have the ability to up-regulate a combination of constitutive mechanical and
1668:
1633:
1621:
684:
648:
562:
516:
59:
2863:
5025:
4510:
4460:
4426:
4375:
4357:
4296:
4211:
4176:
4169:
4144:
4095:
4029:
3565:
3463:
3416:
3352:
3293:
3241:
3177:
3121:
3042:
2985:
2890:
2784:
2656:
2522:
2472:
2454:
2415:
2308:
2250:
2208:
2161:
1673:
616:
508:
496:
213:
3494:
1947:
491:. Pinaceae have distinctive cones with woody scales bearing typically two
4860:
4715:
4614:
4600:
4503:
4469:
4435:
4335:
4328:
4314:
4269:
4251:
4190:
4162:
4116:
4109:
4102:
3982:
3936:
3889:
3840:
3778:
3757:
3642:
1843:
1819:
1788:
are phenolics or polyphenols. These secondary compounds are preserved in
1663:
1469:
1306:
1243:
1082:
914:
850:
781:
764:
750:
739:
651:
are abundant do not seem to have evolved adaptations for bird dispersal.
596:
488:
468:
320:
313:
265:
230:
101:
34:
3224:
2775:
2756:
2648:
2437:
Ran, Jin-Hua; Shen, Ting-Ting; Wang, Ming-Ming; Wang, Xiao-Quan (2018).
4621:
4607:
4524:
4415:
4232:
4218:
4088:
3957:
3943:
3799:
3750:
3696:
3633:
3624:
3571:
3390:
3344:
2939:
2106:
2069:
1793:
1637:
1617:
1390:
1330:
1001:
939:
793:
787:
726:
605:
542:
297:
258:
251:
168:
79:
44:
2506:
2242:
2200:
553:
4865:
4782:
4769:
4649:
4628:
4579:
4559:
4204:
4061:
3860:
3785:
3716:
3687:
3678:
3006:
1855:
1851:
1835:
1735:
1645:
1447:
1060:
705:
679:
609:
586:
582:
281:
84:
4878:
4839:
4677:
3328:
2184:
2098:
2061:
816:
as sister to the pines rather than in the
Laricoidae subfamily with
5020:
4700:
4538:
4483:
4395:
4280:
4183:
4047:
3831:
3500:
2815:
2612:
2595:
2593:
1739:
1715:
1659:
1655:
74:
69:
54:
49:
39:
4774:
3086:
Hudgins, J. W.; Christiansen, E.; Franceschi, V. R. (2004-03-01).
4531:
4321:
4054:
3996:
3904:
3822:
3771:
1789:
1780:
1711:
1641:
1527:
1160:
756:
715:
558:
530:
484:
452:
327:
198:
188:
178:
89:
64:
2903:
2754:
4225:
4130:
3929:
3869:
3655:
3373:
Lewinsohn, Efraim; Gijzen, Mark; Croteau, Rodney (1991-05-01).
2839:"Hemisphere-scale differences in conifer evolutionary dynamics"
1797:
1265:
872:
775:
719:
663:
566:
480:
456:
244:
3522:, covers Californian species and much of western North America
424:
418:
4994:
4761:
4586:
4342:
4010:
4003:
3989:
3964:
3950:
3792:
3732:
3705:
3313:
3085:
1570:
1544:
1486:
1407:
1347:
1178:
1135:
1100:
1019:
957:
799:
769:
744:
731:
710:
601:
590:
578:
492:
472:
464:
334:
304:
288:
272:
237:
142:
2488:
2182:
675:) with the light brown umbo visible on the green cone scales
581:) growing from 2 to 100 metres (7 to 300 feet) tall, mostly
1929:
620:
574:
476:
439:
433:
412:
3488:
2672:
2634:
2802:
from the Early Cretaceous of Yorkshire, United Kingdom".
2373:
2263:
460:
403:
2484:
2482:
1854:. Oleoresin had been found to be a valuable part of the
623:
on each scale. The male cones are small, 0.5–6 cm (
2544:
Domogatskaya, Ksenia V.; Herman, Alexei B. (May 2019).
2443:
Proceedings of the Royal Society B: Biological Sciences
1640:
relatives have been reported from as early as the Late
3432:
3372:
2836:
2479:
507:
opinion) in 11 genera, and the second-largest (after
436:
430:
421:
415:
409:
3501:
Pinaceae on the web page of the Tree-of-Life project
3258:
Nagy, N. E.; Krokene, P.; Solheim, H. (2006-02-01).
3142:
Krokene, P.; Nagy, N. E.; Solheim, H. (2008-01-01).
1869:
400:
406:
3257:
3141:
2543:
5076:
2755:Ashley A. Klymiuk & Ruth A. Stockey (2012).
2030:
2843:Proceedings of the National Academy of Sciences
2436:
1714:, its distribution was limited to the northern
2727:
1874:The topic of defense mechanisms within family
1846:attacks. One well researched resin present in
511:) in geographical range, found in most of the
3551:
3491:French Arboretum of conifers around the world
2957:
2119:
1624:. This is known as the "gnepine" hypothesis.
1974:World Checklist and Bibliography of Conifers
1971:
1814:A common induced defense mechanism used by
1704:appear to be members of Abietoideae, while
3558:
3544:
100:
3632:
3406:
3275:
3223:
3213:
3159:
3103:
3067:). American Journal of Botany 87:314-326.
3024:
2975:
2929:
2880:
2862:
2774:
2611:
2462:
2290:
1636:~313 million years ago. Various possible
830:Ran et al. 2018 & Leslie et al. 2018
3686:
3195:
1967:
1965:
1771:
678:
662:
552:
3912:
2804:International Journal of Plant Sciences
2600:International Journal of Plant Sciences
2330:Stull, Gregory W.; et al. (2021).
2222:Leslie, Andrew B.; et al. (2018).
1672:, known from the Upper Jurassic (lower
1627:
5077:
3428:
3426:
2668:
2666:
2589:
2587:
2221:
483:. The family is included in the order
4682:
4681:
3539:
3368:
3366:
3364:
3362:
3309:
3307:
3305:
3303:
3253:
3251:
3191:
3189:
3187:
3137:
3135:
3133:
3131:
3081:
3079:
3077:
3075:
3073:
3058:
3056:
3054:
3052:
2953:
2951:
2949:
2329:
2126:Molecular Phylogenetics and Evolution
2000:
1962:
1721:
5049:f2275cde-1fb9-474b-92eb-7dc35276f9ab
5000:6D7FD99B-5D8D-F004-BD25-FB45886E1054
4941:f5a67efd-926d-4dab-a08d-3e4d253bce8c
2964:Journal of Integrative Plant Biology
2332:"main.dated.supermatrix.tree.T9.tre"
3506:40 Pine Trees From Around the World
3423:
2958:Zulak, K. G.; Bohlmann, J. (2010).
2679:Journal of Systematic Palaeontology
2663:
2584:
2001:Earle, Christopher J., ed. (2018).
1803:
1658:, with abundant records during the
573:Members of the family Pinaceae are
13:
5013:urn:lsid:ipni.org:names:30000468-2
3526:Pinaceae in Flora of North America
3520:University of California, Berkeley
3359:
3300:
3248:
3184:
3128:
3070:
3049:
3000:
2946:
2897:
14:
5101:
3482:
1870:Active research: methyl jasmonate
1648:) The extinct conifer cone genus
658:
3026:10.1111/j.1469-8137.2005.01436.x
2977:10.1111/j.1744-7909.2010.00910.x
2931:10.1046/j.1365-2745.2002.00715.x
1904:
1892:
812:A revised 2018 phylogeny places
667:An immature second-year cone of
396:
129:
32:
2830:
2791:
2748:
2721:
2628:
2537:
2430:
2367:
2340:10.6084/m9.figshare.14547354.v1
2323:
1680:. Members of the modern genera
2257:
2224:"ajb21143-sup-0004-AppendixS4"
2215:
2176:
2113:
2076:
2024:
1994:
1923:
1730:. Common external stress that
548:
537:are found in the mountains of
357:von Berchtold & Presl 1820
1:
3531:Pinus in USDA Plants Database
2699:10.1080/14772019.2016.1143885
2570:10.1016/j.cretres.2019.01.012
1916:
1899:Close up of bishop pine cones
541:, Mexico, central Japan, and
1768:are both found in the bark.
807:
7:
3196:Sampedro, L. (2014-09-01).
2146:10.1016/j.ympev.2018.08.011
1754:to further their defenses.
1662:across Eurasia. The oldest
10:
5106:
3489:Arboretum de Villardebelle
3317:American Journal of Botany
2762:American Journal of Botany
2637:American Journal of Botany
2495:American Journal of Botany
2400:10.1038/s41477-021-00964-4
2301:10.1038/s41477-021-00964-4
2231:American Journal of Botany
2189:American Journal of Botany
1978:Royal Botanic Gardens, Kew
1858:defense mechanism against
557:Cultivated pine forest in
529:, grows just south of the
369:Delevoryas & Hope 1973
4690:
4569:
4493:
4459:
4425:
4405:
4385:
4365:
4356:
4306:
4288:
4279:
4259:
4250:
4154:
4080:
4071:
4037:
4028:
3974:
3921:
3903:
3879:
3859:
3839:
3830:
3821:
3742:
3724:
3715:
3695:
3677:
3650:
3641:
3623:
3619:
3579:
3277:10.1093/treephys/26.2.159
3105:10.1093/treephys/24.3.251
2292:10.1101/2021.03.13.435279
2031:Craig W. Benkman (1995).
1567:
1541:
1524:
1517:
1510:
1483:
1466:
1459:
1451:
1404:
1387:
1380:
1344:
1327:
1320:
1303:
1296:
1288:
1262:
1254:
1247:
1237:
1175:
1157:
1150:
1132:
1125:
1097:
1079:
1072:
1064:
1016:
998:
991:
954:
936:
929:
911:
904:
896:
869:
861:
854:
844:
832:
829:
533:in Southeast Asia. Major
352:
345:
226:
221:
126:Scientific classification
124:
116:(central foreground) and
108:
99:
23:
3161:10.1093/treephys/28.1.29
3448:10.1023/A:1020714403780
3436:Plant Molecular Biology
3215:10.1093/treephys/tpu082
2864:10.1073/pnas.1213621109
2011:The Gymnosperm Database
495:, and are supported as
2455:10.1098/rspb.2018.1012
2355:Cite journal requires
692:
676:
570:
4962:Paleobiology Database
1972:Aljos Farjon (1998).
1948:10.1139/cjb-2016-0064
1777:Constitutive defenses
1772:Constitutive defenses
1086:(Douglas-firs 5 sp.)
918:(golden larch 1 sp.)
682:
666:
556:
3516:The Jepson Herbarium
1862:. They are found in
1628:Evolutionary history
683:An immature cone of
535:centres of diversity
487:, formerly known as
3495:Gymnosperm Database
2922:2002JEcol..90..839C
2855:2012PNAS..10916217L
2849:(40): 16217–16221.
2776:10.3732/ajb.1100568
2691:2017JSPal..15...69S
2649:10.3732/ajb.1100595
2562:2019CrRes..97...73D
2550:Cretaceous Research
2392:2021NatPl...7.1015S
2283:2021NatPl...7.1015S
2138:2018MolPE.129..106R
2054:1995Oikos..73..221B
1752:chemical strategies
1139:(spruces c 35 sp.)
669:European black pine
513:Northern Hemisphere
120:(right foreground)
3566:Classification of
3391:10.1104/pp.96.1.44
2910:Journal of Ecology
2730:Acta Paleobotanica
2449:(1881): 20181012.
1911:Knobcone pine cone
1834:) and nonvolatile
1722:Defense mechanisms
1182:(pines c.115 sp.)
833:Stull et al. 2021
693:
677:
571:
16:Family of conifers
5072:
5071:
4949:Open Tree of Life
4684:Taxon identifiers
4675:
4674:
4671:
4670:
4667:
4666:
4663:
4662:
4659:
4658:
4367:Cunninghamioideae
4352:
4351:
4246:
4245:
4242:
4241:
4024:
4023:
4020:
4019:
3899:
3898:
3817:
3816:
3813:
3812:
3809:
3808:
3673:
3672:
3669:
3668:
3665:
3664:
2507:10.1002/ajb2.1143
2243:10.1002/ajb2.1143
2201:10.1002/ajb2.1143
2086:Systematic Botany
1987:978-1-900347-54-9
1864:secretory tissues
1728:forest ecosystems
1614:
1613:
1609:
1608:
1600:
1599:
1591:
1590:
1582:
1581:
1556:
1555:
1498:
1497:
1437:
1436:
1428:
1427:
1419:
1418:
1368:
1367:
1359:
1358:
1277:
1276:
1227:
1226:
1218:
1217:
1209:
1208:
1200:
1199:
1191:
1190:
1113:
1112:
1104:(larches 14 sp.)
1050:
1049:
1041:
1040:
1032:
1031:
979:
978:
970:
969:
961:(hemlock 9 sp.)
885:
884:
388:
387:
382:
381:Goroschankin 1904
376:
370:
367:Compsostrobaceae
364:
358:
217:
5097:
5090:Pinales families
5065:
5064:
5052:
5051:
5039:
5038:
5029:
5028:
5016:
5015:
5003:
5002:
4990:
4989:
4980:
4979:
4974:Panartic Flora:
4970:
4969:
4957:
4956:
4944:
4943:
4934:
4933:
4921:
4920:
4918:NBNSYS0000160716
4908:
4907:
4895:
4894:
4882:
4881:
4869:
4868:
4856:
4855:
4843:
4842:
4830:
4829:
4817:
4816:
4804:
4803:
4791:
4790:
4778:
4777:
4765:
4764:
4752:
4751:
4739:
4738:
4726:
4725:
4724:
4711:
4710:
4709:
4679:
4678:
4407:Athrotaxidoideae
4363:
4362:
4286:
4285:
4257:
4256:
4081:Phyllocladoideae
4078:
4077:
4035:
4034:
3919:
3918:
3910:
3909:
3837:
3836:
3828:
3827:
3722:
3721:
3693:
3692:
3684:
3683:
3648:
3647:
3639:
3638:
3630:
3629:
3621:
3620:
3606:Acrogymnospermae
3568:Acrogymnospermae
3560:
3553:
3546:
3537:
3536:
3476:
3475:
3430:
3421:
3420:
3410:
3379:Plant Physiology
3370:
3357:
3356:
3311:
3298:
3297:
3279:
3255:
3246:
3245:
3227:
3217:
3193:
3182:
3181:
3163:
3139:
3126:
3125:
3107:
3083:
3068:
3060:
3047:
3046:
3028:
3004:
2998:
2997:
2979:
2955:
2944:
2943:
2933:
2901:
2895:
2894:
2884:
2866:
2834:
2828:
2827:
2795:
2789:
2788:
2778:
2769:(6): 1069–1082.
2752:
2746:
2745:
2725:
2719:
2718:
2670:
2661:
2660:
2632:
2626:
2625:
2615:
2591:
2582:
2581:
2541:
2535:
2534:
2501:(9): 1531–1544.
2486:
2477:
2476:
2466:
2434:
2428:
2427:
2386:(8): 1015–1025.
2371:
2365:
2364:
2358:
2353:
2351:
2343:
2327:
2321:
2320:
2294:
2277:(8): 1015–1025.
2261:
2255:
2254:
2237:(9): 1531–1544.
2228:
2219:
2213:
2212:
2195:(9): 1531–1544.
2180:
2174:
2173:
2117:
2111:
2110:
2080:
2074:
2073:
2037:
2028:
2022:
2021:
2019:
2017:
1998:
1992:
1991:
1969:
1960:
1959:
1927:
1908:
1896:
1880:methyl jasmonate
1804:Induced defenses
1798:secondary phloem
1794:parenchyma cells
1792:of polyphenolic
1766:induced defenses
1678:Early Cretaceous
1651:Schizolepidopsis
1520:
1519:
1513:
1512:
1462:
1461:
1454:
1453:
1383:
1382:
1323:
1322:
1299:
1298:
1291:
1290:
1257:
1256:
1250:
1249:
1240:
1239:
1233:
1232:
1153:
1152:
1128:
1127:
1075:
1074:
1067:
1066:
1023:(firs c.50 sp.)
994:
993:
932:
931:
907:
906:
899:
898:
864:
863:
857:
856:
847:
846:
840:
839:
827:
826:
646:
645:
641:
638:
632:
631:
627:
446:
445:
442:
441:
438:
435:
432:
427:
426:
423:
420:
417:
414:
411:
408:
405:
402:
380:
374:
368:
362:
356:
212:
134:
133:
104:
94:
31:
27:Temporal range:
21:
20:
5105:
5104:
5100:
5099:
5098:
5096:
5095:
5094:
5075:
5074:
5073:
5068:
5060:
5055:
5047:
5042:
5034:
5032:
5024:
5019:
5011:
5006:
4998:
4993:
4985:
4983:
4975:
4973:
4965:
4960:
4952:
4947:
4939:
4937:
4929:
4924:
4916:
4911:
4903:
4898:
4890:
4885:
4877:
4872:
4864:
4859:
4851:
4846:
4838:
4833:
4825:
4820:
4812:
4807:
4799:
4794:
4786:
4781:
4773:
4768:
4760:
4755:
4747:
4742:
4734:
4729:
4720:
4719:
4714:
4705:
4704:
4699:
4686:
4676:
4655:
4565:
4553:Neocallitropsis
4489:
4455:
4421:
4401:
4381:
4348:
4302:
4275:
4261:Sciadopityaceae
4238:
4150:
4067:
4016:
3970:
3895:
3881:Welwitschiaceae
3875:
3855:
3805:
3738:
3711:
3661:
3615:
3596:Marchantiophyta
3575:
3564:
3485:
3480:
3479:
3431:
3424:
3371:
3360:
3329:10.2307/2656626
3312:
3301:
3264:Tree Physiology
3256:
3249:
3202:Tree Physiology
3194:
3185:
3148:Tree Physiology
3140:
3129:
3092:Tree Physiology
3084:
3071:
3061:
3050:
3013:New Phytologist
3005:
3001:
2956:
2947:
2902:
2898:
2835:
2831:
2796:
2792:
2753:
2749:
2726:
2722:
2671:
2664:
2633:
2629:
2592:
2585:
2542:
2538:
2487:
2480:
2435:
2431:
2372:
2368:
2356:
2354:
2345:
2344:
2328:
2324:
2262:
2258:
2226:
2220:
2216:
2181:
2177:
2118:
2114:
2099:10.2307/2419217
2081:
2077:
2062:10.2307/3545911
2035:
2029:
2025:
2015:
2013:
1999:
1995:
1988:
1970:
1963:
1928:
1924:
1919:
1912:
1909:
1900:
1897:
1872:
1841:
1833:
1825:
1809:Induced defense
1806:
1774:
1734:experience are
1724:
1695:Pseudoaraucaria
1630:
1610:
1601:
1592:
1583:
1557:
1499:
1438:
1429:
1420:
1369:
1360:
1294:Pseudolariceae
1278:
1228:
1219:
1210:
1201:
1192:
1114:
1051:
1042:
1033:
980:
971:
902:Pseudolariceae
886:
876:(cedars 4 sp.)
810:
661:
643:
639:
636:
634:
629:
625:
624:
551:
539:southwest China
523:. One species,
521:montane forests
519:, coastal, and
429:
399:
395:
211:
128:
95:
93:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
37:
30:Jurassic–Recent
29:
28:
25:
17:
12:
11:
5:
5103:
5093:
5092:
5087:
5070:
5069:
5067:
5066:
5062:wfo-7000000470
5053:
5040:
5030:
5017:
5004:
4991:
4981:
4971:
4958:
4945:
4935:
4922:
4909:
4896:
4883:
4870:
4857:
4844:
4831:
4818:
4805:
4792:
4779:
4766:
4753:
4740:
4727:
4712:
4696:
4694:
4688:
4687:
4673:
4672:
4669:
4668:
4665:
4664:
4661:
4660:
4657:
4656:
4654:
4653:
4646:
4643:Hesperocyparis
4639:
4632:
4625:
4618:
4611:
4604:
4597:
4590:
4583:
4575:
4573:
4567:
4566:
4564:
4563:
4556:
4549:
4542:
4535:
4528:
4521:
4518:Pilgerodendron
4514:
4507:
4499:
4497:
4491:
4490:
4488:
4487:
4480:
4473:
4465:
4463:
4457:
4456:
4454:
4453:
4446:
4443:Sequoiadendron
4439:
4431:
4429:
4423:
4422:
4420:
4419:
4411:
4409:
4403:
4402:
4400:
4399:
4391:
4389:
4383:
4382:
4380:
4379:
4371:
4369:
4360:
4354:
4353:
4350:
4349:
4347:
4346:
4339:
4332:
4325:
4318:
4310:
4308:
4304:
4303:
4301:
4300:
4292:
4290:
4283:
4277:
4276:
4274:
4273:
4265:
4263:
4254:
4248:
4247:
4244:
4243:
4240:
4239:
4237:
4236:
4229:
4222:
4215:
4208:
4201:
4194:
4187:
4180:
4173:
4166:
4158:
4156:
4155:Podocarpoideae
4152:
4151:
4149:
4148:
4141:
4134:
4127:
4120:
4113:
4106:
4099:
4092:
4084:
4082:
4075:
4069:
4068:
4066:
4065:
4058:
4051:
4043:
4041:
4032:
4026:
4025:
4022:
4021:
4018:
4017:
4015:
4014:
4007:
4000:
3993:
3986:
3978:
3976:
3972:
3971:
3969:
3968:
3961:
3954:
3947:
3940:
3933:
3925:
3923:
3916:
3907:
3901:
3900:
3897:
3896:
3894:
3893:
3885:
3883:
3877:
3876:
3874:
3873:
3865:
3863:
3857:
3856:
3854:
3853:
3845:
3843:
3834:
3825:
3819:
3818:
3815:
3814:
3811:
3810:
3807:
3806:
3804:
3803:
3796:
3789:
3782:
3775:
3768:
3761:
3754:
3746:
3744:
3740:
3739:
3737:
3736:
3728:
3726:
3719:
3713:
3712:
3710:
3709:
3701:
3699:
3690:
3681:
3675:
3674:
3671:
3670:
3667:
3666:
3663:
3662:
3660:
3659:
3651:
3645:
3636:
3627:
3617:
3616:
3614:
3613:
3608:
3603:
3601:Polypodiophyta
3598:
3593:
3588:
3580:
3577:
3576:
3563:
3562:
3555:
3548:
3540:
3534:
3533:
3528:
3523:
3509:
3503:
3498:
3492:
3484:
3483:External links
3481:
3478:
3477:
3442:(1): 119–133.
3422:
3358:
3323:(3): 302–313.
3299:
3270:(2): 159–167.
3247:
3208:(9): 915–918.
3183:
3127:
3098:(3): 251–264.
3069:
3048:
3019:(2): 353–376.
2999:
2945:
2916:(5): 839–850.
2896:
2829:
2816:10.1086/667228
2810:(8): 917–932.
2790:
2747:
2720:
2662:
2643:(4): 708–720.
2627:
2613:10.1086/714281
2606:(6): 490–507.
2583:
2536:
2478:
2429:
2366:
2357:|journal=
2322:
2256:
2214:
2175:
2112:
2075:
2048:(2): 221–224.
2023:
2005:Pinus merkusii
1993:
1986:
1961:
1942:(9): 863–884.
1921:
1920:
1918:
1915:
1914:
1913:
1910:
1903:
1901:
1898:
1891:
1871:
1868:
1860:biotic attacks
1839:
1838:resin acids (C
1831:
1828:sesquiterpenes
1823:
1805:
1802:
1773:
1770:
1723:
1720:
1629:
1626:
1612:
1611:
1607:
1606:
1603:
1602:
1598:
1597:
1594:
1593:
1589:
1588:
1585:
1584:
1580:
1579:
1576:
1575:
1566:
1563:
1562:
1559:
1558:
1554:
1553:
1550:
1549:
1540:
1537:
1536:
1533:
1532:
1523:
1518:
1516:
1511:
1509:
1505:
1504:
1501:
1500:
1496:
1495:
1492:
1491:
1482:
1479:
1478:
1475:
1474:
1465:
1460:
1458:
1452:
1450:
1444:
1443:
1440:
1439:
1435:
1434:
1431:
1430:
1426:
1425:
1422:
1421:
1417:
1416:
1413:
1412:
1403:
1400:
1399:
1396:
1395:
1386:
1381:
1379:
1375:
1374:
1371:
1370:
1366:
1365:
1362:
1361:
1357:
1356:
1353:
1352:
1343:
1340:
1339:
1336:
1335:
1326:
1321:
1319:
1316:
1315:
1312:
1311:
1302:
1297:
1295:
1289:
1287:
1284:
1283:
1280:
1279:
1275:
1274:
1271:
1270:
1261:
1255:
1253:
1248:
1246:
1238:
1236:
1231:
1229:
1225:
1224:
1221:
1220:
1216:
1215:
1212:
1211:
1207:
1206:
1203:
1202:
1198:
1197:
1194:
1193:
1189:
1188:
1185:
1184:
1174:
1171:
1170:
1167:
1166:
1156:
1151:
1149:
1146:
1145:
1142:
1141:
1131:
1126:
1124:
1120:
1119:
1116:
1115:
1111:
1110:
1107:
1106:
1096:
1093:
1092:
1089:
1088:
1078:
1073:
1071:
1065:
1063:
1057:
1056:
1053:
1052:
1048:
1047:
1044:
1043:
1039:
1038:
1035:
1034:
1030:
1029:
1026:
1025:
1015:
1012:
1011:
1008:
1007:
997:
992:
990:
986:
985:
982:
981:
977:
976:
973:
972:
968:
967:
964:
963:
953:
950:
949:
946:
945:
935:
930:
928:
925:
924:
921:
920:
910:
905:
903:
897:
895:
892:
891:
888:
887:
883:
882:
879:
878:
868:
862:
860:
855:
853:
845:
843:
838:
835:
834:
831:
809:
806:
805:
804:
761:
736:
723:
691:) with no umbo
660:
659:Classification
657:
649:tree squirrels
550:
547:
526:Pinus merkusii
386:
385:
384:
383:
377:
373:†Kranneraceae
371:
365:
359:
350:
349:
343:
342:
341:
340:
339:
338:
331:
324:
310:
309:
308:
294:
293:
292:
278:
277:
276:
269:
262:
255:
248:
241:
224:
223:
219:
218:
206:
202:
201:
196:
192:
191:
186:
182:
181:
176:
172:
171:
166:
159:
158:
153:
146:
145:
140:
136:
135:
122:
121:
106:
105:
97:
96:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
33:
26:
15:
9:
6:
4:
3:
2:
5102:
5091:
5088:
5086:
5083:
5082:
5080:
5063:
5058:
5054:
5050:
5045:
5041:
5037:
5031:
5027:
5022:
5018:
5014:
5009:
5005:
5001:
4996:
4992:
4988:
4982:
4978:
4972:
4968:
4963:
4959:
4955:
4950:
4946:
4942:
4936:
4932:
4927:
4923:
4919:
4914:
4910:
4906:
4901:
4897:
4893:
4888:
4884:
4880:
4875:
4871:
4867:
4862:
4858:
4854:
4849:
4845:
4841:
4836:
4832:
4828:
4823:
4819:
4815:
4810:
4806:
4802:
4797:
4793:
4789:
4784:
4780:
4776:
4771:
4767:
4763:
4758:
4754:
4750:
4745:
4741:
4737:
4732:
4728:
4723:
4717:
4713:
4708:
4702:
4698:
4697:
4695:
4693:
4689:
4685:
4680:
4652:
4651:
4647:
4645:
4644:
4640:
4638:
4637:
4636:Xanthocyparis
4633:
4631:
4630:
4626:
4624:
4623:
4619:
4617:
4616:
4612:
4610:
4609:
4605:
4603:
4602:
4598:
4596:
4595:
4594:Chamaecyparis
4591:
4589:
4588:
4584:
4582:
4581:
4577:
4576:
4574:
4572:
4571:Cupressoideae
4568:
4562:
4561:
4557:
4555:
4554:
4550:
4548:
4547:
4546:Widdringtonia
4543:
4541:
4540:
4536:
4534:
4533:
4529:
4527:
4526:
4522:
4520:
4519:
4515:
4513:
4512:
4508:
4506:
4505:
4501:
4500:
4498:
4496:
4495:Callitroideae
4492:
4486:
4485:
4481:
4479:
4478:
4477:Glyptostrobus
4474:
4472:
4471:
4467:
4466:
4464:
4462:
4458:
4452:
4451:
4447:
4445:
4444:
4440:
4438:
4437:
4433:
4432:
4430:
4428:
4424:
4418:
4417:
4413:
4412:
4410:
4408:
4404:
4398:
4397:
4393:
4392:
4390:
4388:
4387:Taiwanioideae
4384:
4378:
4377:
4373:
4372:
4370:
4368:
4364:
4361:
4359:
4355:
4345:
4344:
4340:
4338:
4337:
4333:
4331:
4330:
4326:
4324:
4323:
4319:
4317:
4316:
4312:
4311:
4309:
4305:
4299:
4298:
4294:
4293:
4291:
4289:Cephalotaxeae
4287:
4284:
4282:
4278:
4272:
4271:
4267:
4266:
4264:
4262:
4258:
4255:
4253:
4249:
4235:
4234:
4230:
4228:
4227:
4223:
4221:
4220:
4216:
4214:
4213:
4209:
4207:
4206:
4202:
4200:
4199:
4198:Falcatifolium
4195:
4193:
4192:
4188:
4186:
4185:
4181:
4179:
4178:
4174:
4172:
4171:
4167:
4165:
4164:
4160:
4159:
4157:
4153:
4147:
4146:
4142:
4140:
4139:
4138:Lepidothamnus
4135:
4133:
4132:
4128:
4126:
4125:
4124:Lagarostrobos
4121:
4119:
4118:
4114:
4112:
4111:
4107:
4105:
4104:
4100:
4098:
4097:
4093:
4091:
4090:
4086:
4085:
4083:
4079:
4076:
4074:
4073:Podocarpaceae
4070:
4064:
4063:
4059:
4057:
4056:
4052:
4050:
4049:
4045:
4044:
4042:
4040:
4039:Araucariaceae
4036:
4033:
4031:
4027:
4013:
4012:
4008:
4006:
4005:
4001:
3999:
3998:
3994:
3992:
3991:
3987:
3985:
3984:
3980:
3979:
3977:
3973:
3967:
3966:
3962:
3960:
3959:
3955:
3953:
3952:
3948:
3946:
3945:
3941:
3939:
3938:
3934:
3932:
3931:
3927:
3926:
3924:
3920:
3917:
3915:
3911:
3908:
3906:
3902:
3892:
3891:
3887:
3886:
3884:
3882:
3878:
3872:
3871:
3867:
3866:
3864:
3862:
3858:
3852:
3851:
3847:
3846:
3844:
3842:
3838:
3835:
3833:
3829:
3826:
3824:
3820:
3802:
3801:
3797:
3795:
3794:
3790:
3788:
3787:
3783:
3781:
3780:
3776:
3774:
3773:
3769:
3767:
3766:
3765:Encephalartos
3762:
3760:
3759:
3755:
3753:
3752:
3748:
3747:
3745:
3741:
3735:
3734:
3730:
3729:
3727:
3723:
3720:
3718:
3714:
3708:
3707:
3703:
3702:
3700:
3698:
3694:
3691:
3689:
3685:
3682:
3680:
3676:
3658:
3657:
3653:
3652:
3649:
3646:
3644:
3640:
3637:
3635:
3631:
3628:
3626:
3622:
3618:
3612:
3609:
3607:
3604:
3602:
3599:
3597:
3594:
3592:
3589:
3586:
3582:
3581:
3578:
3573:
3569:
3561:
3556:
3554:
3549:
3547:
3542:
3541:
3538:
3532:
3529:
3527:
3524:
3521:
3517:
3513:
3512:Jepson eFlora
3510:
3508:by The Spruce
3507:
3504:
3502:
3499:
3496:
3493:
3490:
3487:
3486:
3473:
3469:
3465:
3461:
3457:
3453:
3449:
3445:
3441:
3437:
3429:
3427:
3418:
3414:
3409:
3404:
3400:
3396:
3392:
3388:
3384:
3380:
3376:
3369:
3367:
3365:
3363:
3354:
3350:
3346:
3342:
3338:
3334:
3330:
3326:
3322:
3318:
3310:
3308:
3306:
3304:
3295:
3291:
3287:
3283:
3278:
3273:
3269:
3265:
3261:
3254:
3252:
3243:
3239:
3235:
3231:
3226:
3221:
3216:
3211:
3207:
3203:
3199:
3192:
3190:
3188:
3179:
3175:
3171:
3167:
3162:
3157:
3153:
3149:
3145:
3138:
3136:
3134:
3132:
3123:
3119:
3115:
3111:
3106:
3101:
3097:
3093:
3089:
3082:
3080:
3078:
3076:
3074:
3066:
3059:
3057:
3055:
3053:
3044:
3040:
3036:
3032:
3027:
3022:
3018:
3014:
3010:
3003:
2995:
2991:
2987:
2983:
2978:
2973:
2969:
2965:
2961:
2954:
2952:
2950:
2941:
2937:
2932:
2927:
2923:
2919:
2915:
2911:
2907:
2900:
2892:
2888:
2883:
2878:
2874:
2870:
2865:
2860:
2856:
2852:
2848:
2844:
2840:
2833:
2825:
2821:
2817:
2813:
2809:
2805:
2801:
2794:
2786:
2782:
2777:
2772:
2768:
2764:
2763:
2758:
2751:
2743:
2739:
2735:
2731:
2724:
2716:
2712:
2708:
2704:
2700:
2696:
2692:
2688:
2684:
2680:
2676:
2669:
2667:
2658:
2654:
2650:
2646:
2642:
2638:
2631:
2623:
2619:
2614:
2609:
2605:
2601:
2597:
2590:
2588:
2579:
2575:
2571:
2567:
2563:
2559:
2555:
2551:
2547:
2540:
2532:
2528:
2524:
2520:
2516:
2512:
2508:
2504:
2500:
2496:
2492:
2485:
2483:
2474:
2470:
2465:
2460:
2456:
2452:
2448:
2444:
2440:
2433:
2425:
2421:
2417:
2413:
2409:
2405:
2401:
2397:
2393:
2389:
2385:
2381:
2380:Nature Plants
2377:
2370:
2362:
2349:
2341:
2337:
2333:
2326:
2318:
2314:
2310:
2306:
2302:
2298:
2293:
2288:
2284:
2280:
2276:
2272:
2271:Nature Plants
2268:
2260:
2252:
2248:
2244:
2240:
2236:
2232:
2225:
2218:
2210:
2206:
2202:
2198:
2194:
2190:
2186:
2179:
2171:
2167:
2163:
2159:
2155:
2151:
2147:
2143:
2139:
2135:
2131:
2127:
2123:
2116:
2108:
2104:
2100:
2096:
2092:
2088:
2087:
2079:
2071:
2067:
2063:
2059:
2055:
2051:
2047:
2043:
2042:
2034:
2027:
2012:
2008:
2006:
1997:
1989:
1983:
1979:
1975:
1968:
1966:
1957:
1953:
1949:
1945:
1941:
1937:
1933:
1926:
1922:
1907:
1902:
1895:
1890:
1889:
1888:
1885:
1881:
1877:
1867:
1865:
1861:
1857:
1853:
1849:
1845:
1837:
1829:
1821:
1817:
1812:
1810:
1801:
1799:
1795:
1791:
1787:
1782:
1778:
1769:
1767:
1763:
1759:
1755:
1753:
1749:
1745:
1741:
1737:
1733:
1729:
1719:
1717:
1713:
1709:
1708:
1703:
1702:
1697:
1696:
1691:
1688:(spruce) and
1687:
1683:
1679:
1675:
1671:
1670:
1669:Eathiestrobus
1665:
1661:
1657:
1653:
1652:
1647:
1643:
1639:
1635:
1634:Carboniferous
1625:
1623:
1622:Carboniferous
1619:
1605:
1604:
1596:
1595:
1587:
1586:
1578:
1577:
1574:
1573:
1572:
1565:
1564:
1561:
1560:
1552:
1551:
1548:
1547:
1546:
1539:
1538:
1535:
1534:
1531:
1530:
1529:
1522:
1521:
1515:
1514:
1507:
1506:
1503:
1502:
1494:
1493:
1490:
1489:
1488:
1481:
1480:
1477:
1476:
1473:
1472:
1471:
1464:
1463:
1456:
1455:
1449:
1446:
1445:
1442:
1441:
1433:
1432:
1424:
1423:
1415:
1414:
1411:
1410:
1409:
1402:
1401:
1398:
1397:
1394:
1393:
1392:
1385:
1384:
1377:
1376:
1373:
1372:
1364:
1363:
1355:
1354:
1351:
1350:
1349:
1342:
1341:
1338:
1337:
1334:
1333:
1332:
1325:
1324:
1318:
1317:
1314:
1313:
1310:
1309:
1308:
1301:
1300:
1293:
1292:
1286:
1285:
1282:
1281:
1273:
1272:
1269:
1268:
1267:
1259:
1258:
1252:
1251:
1245:
1242:
1241:
1235:
1234:
1230:
1223:
1222:
1214:
1213:
1205:
1204:
1196:
1195:
1187:
1186:
1183:
1181:
1180:
1173:
1172:
1169:
1168:
1165:
1163:
1162:
1155:
1154:
1148:
1147:
1144:
1143:
1140:
1138:
1137:
1130:
1129:
1122:
1121:
1118:
1117:
1109:
1108:
1105:
1103:
1102:
1095:
1094:
1091:
1090:
1087:
1085:
1084:
1077:
1076:
1069:
1068:
1062:
1059:
1058:
1055:
1054:
1046:
1045:
1037:
1036:
1028:
1027:
1024:
1022:
1021:
1014:
1013:
1010:
1009:
1006:
1004:
1003:
996:
995:
988:
987:
984:
983:
975:
974:
966:
965:
962:
960:
959:
952:
951:
948:
947:
944:
942:
941:
934:
933:
927:
926:
923:
922:
919:
917:
916:
909:
908:
901:
900:
894:
893:
890:
889:
881:
880:
877:
875:
874:
866:
865:
859:
858:
852:
849:
848:
842:
841:
837:
836:
828:
825:
823:
819:
815:
802:
801:
796:
795:
790:
789:
784:
783:
778:
777:
772:
771:
766:
762:
759:
758:
753:
752:
747:
746:
741:
737:
734:
733:
728:
724:
721:
717:
713:
712:
707:
703:
702:
701:
697:
690:
686:
685:Norway spruce
681:
674:
670:
665:
656:
652:
650:
622:
618:
613:
611:
607:
603:
599:
598:
593:
592:
588:
584:
580:
576:
568:
564:
563:Western Ghats
560:
555:
546:
544:
540:
536:
532:
528:
527:
522:
518:
514:
510:
506:
502:
501:morphological
498:
494:
490:
486:
482:
478:
474:
470:
466:
462:
458:
454:
450:
444:
393:
378:
372:
366:
360:
354:
353:
351:
348:
344:
337:
336:
332:
330:
329:
325:
323:
322:
318:
317:
316:
315:
311:
307:
306:
302:
301:
300:
299:
295:
291:
290:
286:
285:
284:
283:
279:
275:
274:
270:
268:
267:
263:
261:
260:
256:
254:
253:
249:
247:
246:
242:
240:
239:
235:
234:
233:
232:
228:
227:
225:
220:
215:
210:
207:
204:
203:
200:
197:
194:
193:
190:
187:
184:
183:
180:
177:
174:
173:
170:
167:
164:
161:
160:
157:
156:Tracheophytes
154:
151:
148:
147:
144:
141:
138:
137:
132:
127:
123:
119:
115:
111:
107:
103:
98:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
22:
19:
4691:
4648:
4641:
4634:
4627:
4620:
4613:
4606:
4599:
4592:
4585:
4578:
4558:
4551:
4544:
4537:
4530:
4523:
4516:
4511:Austrocedrus
4509:
4502:
4482:
4475:
4468:
4461:Taxodioideae
4448:
4441:
4434:
4427:Sequoioideae
4414:
4394:
4376:Cunninghamia
4374:
4358:Cupressaceae
4341:
4334:
4327:
4320:
4313:
4297:Cephalotaxus
4295:
4268:
4231:
4224:
4217:
4212:Retrophyllum
4210:
4203:
4196:
4189:
4182:
4177:Pherosphaera
4175:
4170:Microcachrys
4168:
4161:
4145:Phyllocladus
4143:
4136:
4129:
4122:
4115:
4108:
4101:
4096:Pectinopitys
4094:
4087:
4060:
4053:
4046:
4030:Araucariales
4009:
4002:
3995:
3988:
3981:
3963:
3956:
3949:
3942:
3935:
3928:
3913:
3888:
3868:
3848:
3798:
3791:
3784:
3777:
3770:
3763:
3756:
3749:
3731:
3704:
3654:
3611:Angiospermae
3439:
3435:
3385:(1): 44–49.
3382:
3378:
3320:
3316:
3267:
3263:
3225:10261/105595
3205:
3201:
3154:(1): 29–35.
3151:
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3095:
3091:
3064:
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3002:
2970:(1): 86–97.
2967:
2963:
2913:
2909:
2899:
2846:
2842:
2832:
2807:
2803:
2799:
2793:
2766:
2760:
2750:
2733:
2729:
2723:
2685:(1): 69–81.
2682:
2678:
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2636:
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2603:
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2549:
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2494:
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2442:
2432:
2383:
2379:
2369:
2348:cite journal
2334:. Figshare.
2325:
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2270:
2259:
2234:
2230:
2217:
2192:
2188:
2178:
2129:
2125:
2115:
2093:(1): 91–97.
2090:
2084:
2078:
2045:
2039:
2026:
2014:. Retrieved
2010:
2004:
1996:
1973:
1939:
1935:
1925:
1883:
1875:
1873:
1847:
1815:
1813:
1807:
1796:(PP) in the
1785:
1775:
1762:Constitutive
1757:
1756:
1747:
1743:
1731:
1725:
1707:Pityostrobus
1705:
1701:Obirastrobus
1699:
1693:
1689:
1685:
1681:
1674:Kimmeridgian
1667:
1649:
1631:
1615:
1569:
1568:
1543:
1542:
1526:
1525:
1485:
1484:
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1177:
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1158:
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999:
956:
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813:
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792:
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780:
774:
768:
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749:
743:
730:
709:
698:
694:
688:
672:
653:
614:
595:
589:
585:(except the
572:
524:
509:Cupressaceae
497:monophyletic
448:
391:
389:
333:
326:
319:
312:
303:
296:
287:
280:
271:
264:
257:
250:
243:
236:
229:
208:
169:Gymnospermae
162:
149:
117:
113:
109:
18:
4861:iNaturalist
4716:Wikispecies
4615:Platycladus
4601:Tetraclinis
4504:Papuacedrus
4470:Cryptomeria
4436:Metasequoia
4336:Pseudotaxus
4329:Austrotaxus
4315:Amentotaxus
4270:Sciadopitys
4252:Cupressales
4191:Dacrycarpus
4163:Saxegothaea
4117:Parasitaxus
4110:Prumnopitys
4103:Sundacarpus
3983:Pseudotsuga
3937:Pseudolarix
3922:Abietoideae
3890:Welwitschia
3841:Ephedraceae
3779:Ceratozamia
3758:Lepidozamia
3643:Ginkgoaceae
3583:Supergroup
3572:Gymnosperms
2736:: 379–389.
2132:: 106–116.
1844:bark beetle
1664:crown group
1470:Pseudotsuga
1307:Pseudolarix
1244:Abietoideae
1083:Pseudotsuga
915:Pseudolarix
851:Abietoideae
822:Pseudotsuga
782:Pseudolarix
765:Abietoideae
751:Pseudotsuga
740:Laricoideae
718:(below the
689:Picea abies
673:Pinus nigra
615:The female
597:Pseudolarix
561:, southern
549:Description
489:Coniferales
449:pine family
355:Abietaceae
321:Pseudotsuga
314:Laricoideae
266:Pseudolarix
231:Abietoideae
5079:Categories
4879:30000468-2
4622:Calocedrus
4608:Microbiota
4525:Libocedrus
4416:Athrotaxis
4233:Podocarpus
4219:Afrocarpus
4089:Halocarpus
3958:Keteleeria
3944:Nothotsuga
3800:Microcycas
3751:Macrozamia
3743:Zamioideae
3697:Cycadaceae
3634:Ginkgoales
3625:Ginkgoidae
3497:– Pinaceae
1917:References
1638:stem-group
1618:Gnetophyta
1391:Keteleeria
1331:Nothotsuga
1002:Keteleeria
940:Nothotsuga
794:Nothotsuga
788:Keteleeria
763:Subfamily
738:Subfamily
727:Piceoideae
725:Subfamily
704:Subfamily
610:cotyledons
606:monoecious
543:California
379:Piceaceae
375:Corda 1866
361:Cedraceae
298:Piceoideae
259:Nothotsuga
252:Keteleeria
175:Division:
112:(golden),
4783:FloraBase
4650:Juniperus
4629:Cupressus
4580:Thujopsis
4560:Callitris
4307:Taxoideae
4205:Dacrydium
4062:Araucaria
3975:Pinoideae
3861:Gnetaceae
3786:Stangeria
3725:Diooideae
3717:Zamiaceae
3688:Cycadales
3679:Cycadidae
3591:Bryophyta
3456:0167-4412
3399:0032-0889
3337:1537-2197
3286:0829-318X
3234:0829-318X
3170:0829-318X
3114:0829-318X
3035:1469-8137
2873:0027-8424
2707:1477-2019
2622:1058-5893
2578:134849082
2556:: 73–93.
2515:1537-2197
2424:236141481
2408:2055-0278
2317:232282918
2154:1055-7903
2016:March 17,
1956:1916-2790
1852:oleoresin
1836:diterpene
1781:phenolics
1736:herbivore
1684:(pines),
1646:Lopingian
1457:Lariceae
1448:Pinoideae
1378:Abieteae
1070:Lariceae
1061:Pinoideae
989:Abieteae
808:Phylogeny
706:Pinoideae
587:deciduous
583:evergreen
505:taxonomic
363:Vest 1818
282:Pinoideae
189:Pinopsida
179:Pinophyta
139:Kingdom:
5085:Pinaceae
5044:VicFlora
5033:VASCAN:
5026:42000417
5021:Tropicos
4987:pinaceae
4814:Pinaceae
4722:Pinaceae
4701:Wikidata
4692:Pinaceae
4539:Fitzroya
4484:Taxodium
4396:Taiwania
4281:Taxaceae
4184:Acmopyle
4048:Wollemia
3914:Pinaceae
3832:Gnetales
3570:(living
3472:21153303
3464:12602896
3417:16668184
3353:10718991
3294:16356912
3242:25261122
3178:17938111
3122:14704135
3065:Pinaceae
3043:15998390
2994:26043965
2986:20074143
2891:22988083
2824:85402168
2785:22623610
2742:54653621
2715:88292891
2657:22491001
2531:52120430
2523:30157290
2473:29925623
2416:34282286
2309:34282286
2251:30157290
2209:30157290
2170:52110440
2162:30153503
1884:Pinaceae
1876:Pinaceae
1848:Pinaceae
1816:Pinaceae
1790:vacuoles
1786:Pinaceae
1758:Pinaceae
1748:Pinaceae
1744:Pinaceae
1740:pathogen
1732:Pinaceae
1716:Laurasia
1660:Jurassic
1656:Triassic
1260:Cedreae
1164:(1 sp.)
1005:(3 sp.)
943:(1 sp.)
867:Cedreae
602:resinous
577:(rarely
499:by both
465:hemlocks
392:Pinaceae
347:Synonyms
209:Pinaceae
205:Family:
24:Pinaceae
4707:Q101680
4532:Diselma
4450:Sequoia
4322:Torreya
4055:Agathis
3997:Cathaya
3905:Pinales
3850:Ephedra
3823:Pinidae
3772:Bowenia
3408:1080711
3345:2656626
2940:3072253
2918:Bibcode
2882:3479534
2851:Bibcode
2687:Bibcode
2558:Bibcode
2464:6030518
2388:Bibcode
2287:bioRxiv
2279:Bibcode
2134:Bibcode
2107:2419217
2070:3545911
2050:Bibcode
1856:conifer
1642:Permian
1528:Cathaya
1508:Pineae
1161:Cathaya
1123:Pineae
814:Cathaya
757:Cathaya
716:abaxial
642:⁄
628:⁄
569:, India
559:Vagamon
531:equator
485:Pinales
481:spruces
473:larches
453:conifer
328:Cathaya
222:Genera
214:Lindley
199:Pinales
195:Order:
185:Class:
143:Plantae
4954:994067
4938:NZOR:
4892:114296
4226:Nageia
4131:Manoao
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3870:Gnetum
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1266:Cedrus
873:Cedrus
797:, and
776:Cedrus
754:, and
720:phloem
579:shrubs
567:Kerala
517:boreal
493:ovules
469:piñons
457:cedars
451:, are
447:), or
245:Cedrus
4995:Plazi
4984:PPE:
4967:54794
4905:18030
4887:IRMNG
4866:47562
4827:10691
4809:FoAO2
4801:10691
4788:22737
4775:1PINF
4749:625M7
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4587:Thuja
4343:Taxus
4011:Pinus
4004:Picea
3990:Larix
3965:Abies
3951:Tsuga
3793:Zamia
3733:Dioon
3706:Cycas
3585:Plant
3468:S2CID
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2990:S2CID
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2041:Oikos
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1101:Larix
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958:Tsuga
818:Larix
800:Tsuga
770:Abies
745:Larix
732:Picea
711:Pinus
621:seeds
617:cones
591:Larix
575:trees
477:pines
335:Larix
305:Picea
289:Pinus
273:Tsuga
238:Abies
163:Clade
150:Clade
118:Pinus
114:Abies
110:Larix
5008:POWO
4931:3318
4926:NCBI
4900:ITIS
4874:IPNI
4848:GRIN
4840:3925
4835:GBIF
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