Lodgepole pine produces viable seed at an early age, commonly 5 to 10 years; germination percentage is as high as that of seed borne by mature trees. Pollen flowers have been observed on 2-0 seedlings in the Lucky Peak Nursery near Boise, ID.
Lodgepole pine is a prolific seed producer. Good crops can be expected at 1- to 3-year intervals, with light crops intervening. The cones withstand below freezing temperatures and are not generally affected by cone- and seed-feeding insects. Only squirrels and coreid bugs are significant seed predators. Seed production should not be taken for granted, however. Complete seed crop failures have occurred at 2800 m (9,200 ft) in northwest Wyoming for 2 to 4 years in a row (42).
Cone production of individual dominant and codominant trees can vary from a few hundred to a few thousand per tree (37). Cones are persistent, and serotinous (closed) cones accumulate for decades. Annual production may run from 173,000 to 790,000 seeds per hectare (70,000 to 320,000/acre) with half to one-third available for annual seedfall (27). An annual seedfall of 99,000 to 222,000 seeds per hectare (40,000 to 90,000/acre) was found in central Montana (58). These figures might be considered typical for interior lodgepole pine where some portion of the trees are of the serotinous type. In Oregon, where the nonserotinous cone habit is prevalent, seedfall ranged from about 35,000 to over 1.2 million/ha (14,000 to 500,000/acre) (21). Most years seedfall was on the order of hundreds of thousands per hectare. Where stored seeds are in the millions per hectare (in closed cones), the number of seeds stored is probably 10 times that of seeds produced annually (37).
Although the number of fully developed seeds per cone varies from as few as 1 to 2 to as many as 50, a normal average for large cone lots in the Rocky Mountains is from 10 to 24 seeds per cone (42). Sierra Nevada populations range from 5 to 37 seeds per cone (20).
The serotinous cone habit varies over wide geographic areas as well as locally (37). Serotinous cones are not common in eastern Oregon, rare in coastal populations, and absent in the Sierra Nevada and southern California and Baja California populations (20). Although common in the Rocky Mountains, this cone habit varies considerably (37). Many stands in the Rockies have less than 50 percent serotinous-cone trees.
Lodgepole pine has long been regarded as a fire-maintained subclimax type. Its ability to regenerate in extremely dense stands to the exclusion of other species can be attributed to the closed cone habit. Millions of seeds per hectare are held in reserve for many years and are readily available to germinate on the seedbed prepared by fire. Recent evidence seems to indicate that fire selects strongly for the closed cone habit (49).
Serotinous cones do not open at maturity because of a resinous bond between the cone scales. The bonds break with temperatures between 45° and 60° C (113° to 140° F) (48), and cone scales are then free to open hygroscopically. Large quantities of seeds are thus available for regenerating a stand following fire. Closed cones at or near the soil surface (less than 30 cm or about 12 in) are also subjected to temperatures from insolation sufficient to open them and may provide seed in harvested areas. Some seeds may be damaged by fire, however, particularly in fires burning in logging slash.
Seeds stored in serotinous cones on the tree remain viable for years. Apparently, prolonged viability can be maintained so long as cones or seeds are not in contact with the ground. Once cones are on the ground, cones open. Damping-off fungi may infect the seed, rodents may feed on the seeds, or germination may occur; for the most part, seeds are not stored in the soil.
Lodgepole pine has relatively small seeds for pine. Seed weights vary considerably, ranging from 2.3 mg (0.04 grains) per seed in the Interior of Canada to 11.4 mg (0.18 grains) per seed in the Sierra Nevada (20). Lodgepole pine seeds average about 298,000 cleaned seeds per kilogram (135,000/lb) for varieties contorta, 258,000/kg (117,000/lb) for murrayana, and 207,000/kg (94,000/lb) for latifolia (54). Density of seedfall 20 m (66 ft) from the timber edge is only 10 to 30 percent of that at the timber edge for stands in the Rocky Mountains (fig. 1) (42). Dispersal of sufficient seed to adequately restock an area often is only about 60 m (200 ft) (23,38). Prevailing winds, thermal effects, or scudding on the snow may disperse seeds far beyond these distances, however.
Figure 1- Sound seed per hectare as a function of distance
from the nearest timber edge.
The annual seedfall from nonserotinous cones helps in restocking relatively minor disturbances in the stand, in maintaining the presence of lodgepole pine in mixed stands, and in expanding conifers into other vegetative types. Seldom do we find stands without some trees of the open-coned type. The efficacy of this seed source can be seen in the dense stands of lodgepole pine along road cuts, powerline rights-of-way, and ditches or where disturbance occurs near lodgepole pine stands.
Studies of seedfall have shown variation in the number of seeds released soon after cone maturation, but most seeds (80 to 90 percent) are released before the following growing season (27).
Where large amounts of seed are stored in serotinous cones, a most effective means of seed dispersal in clearcuts is from cones attached to the slash and those knocked from the slash and scattered over the forest floor during slash disposal. Many cones on or near the ground are opened by normal summer soil surface temperatures (35). In Montana 83 percent of the cones on the ground opened the first year on south slopes compared to 40 percent on north slopes. Maximum seed release from serotinous cones near the ground takes place during the first year of exposure. In fact, cones may open after the first few minutes of exposure to temperatures high enough to break the resinous bonds.
In slash, serotinous cones that are well above the ground behave like those on a tree- they remain closed, and stored seeds remain viable for years.
Seeds in unopened cones and those released from the slash may also be lost to rodents, fungi, and other destructive agents. Seeds from closed cones are usually available only for the first growing season following harvest, but stocking from open-cone seed sources can continue to increase for several years.
Slash disposal on areas where regeneration is planned from serotinous cones must be carefully planned and executed. Seed supply will be largely destroyed if slash to be burned is piled before cones have had a chance to open (38). Piling slash should be delayed until sufficient cones have opened to assure adequate stocking. Piling then scatters seeds and opened cones and helps prepare the seedbed. Piling slash after germination can also decrease stocking because young seedlings are trampled or buried.
Broadcast burning may hasten release of seeds from cones not in a position to open from high soil-surface temperatures. Some seeds will be destroyed, however; the amount will vary with fire intensity.