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Seed Production and Dissemination- Trees
reach cone-bearing age when relatively
young: 25 years old and 1.5 to 3.0 m (5
to 10 ft) tall. Seeds are not produced
in quantity, however, until age 75 to
100, but the long-lived pinyons continue
to bear for a few centuries. A mature
pinyon usually has a broad and rounded
or irregular crown, which is often
almost as wide as the tree height. Such
trees are the heaviest seed producers,
since cones are found mostly in the
upper half of the crown near the ends of
branches. Each cone contains about 10 to
20 seeds, which average only 4,190/kg
(1,900/lb) because of their large size.
A large tree in a good crop year may
yield over 9.1 kg (20 lb) of seed, and
better stands will produce an estimated
336 kg/ha (300 lb/acre).
Cone crops are either good or poor,
often with cones practically absent, but
seldom intermediate. Although good crops
tend to be localized and occur at
irregular and infrequent intervals, some
are found nearly every year somewhere
over the widespread range of the
species. Furthermore, cone bearing tends
to be synchronous over large
geographical areas, a condition
considered to be an evolved mechanism to
counteract seed predation. On an
average, substantial crops are produced
every 4 to 7 years, but shorter
intervals of 2 to 5 years elapse with
individual trees or in certain
localities. Generally, crops occur more
frequently on better sites over the
optimum range of pinyon than at the
extreme limits.
The large, wingless seeds of pinyon are
not adapted to wind dissemination.
Instead, seed dispersal beyond tree
crowns depends upon the behavior of four
corvid species of birds- Clark's
nutcracker, Steller's jay, scrub jay,
and pinyon jay (8). Although these
species eat great quantities of seed
during the fall and may be greater
predators than rodents, they also cache
large amounts for consumption during
ensuing winter months. Some of these
buried seeds are not recovered by the
birds, thus providing a seed source for
subsequent germination and seedling
establishment, particularly if caches
are located in a suitable
microenvironment, such as alongside
shrubs or downed trees.
Steller's and scrub jays collect seed
only from open cones. In contrast,
pinyon jays and Clark's nutcrackers
forage from green cones, from which
seeds are deftly extracted, and then
from open cones as the season
progresses. Clark's nutcrackers and
Steller's jays probably contribute
little towards regenerating existing
woodland sites because their caches are
located at higher elevations in
ponderosa pine and mixed conifer forests
or in the ecotone above pinyon-juniper
woodlands. Thus, these species tend to
expand woodlands to upper elevations. In
contrast, scrub jays and pinyon jays
cache seeds in woodland areas, the
former in small, local territories,
whereas the latter transport seeds up to
12 kilometers (7.5 mi). Pinyon jays live
in flocks of 50 to 500 birds, and it has
been estimated that during a substantial
seed year in New Mexico, about 4.5
million seeds were cached by a single
flock. Even scrub jays, which do not
exhibit flock behavior can be important
seed dispersers-a single pair of birds
may harvest and cache about 13,000 seeds
from a particular crop.
Pinyon jays can carry an average of up
to 56 seeds in an expandable esophagus.
Scrub jays lack this adaption, and the
amount of seed that can be transported
at one time is limited to 5 or fewer
seeds held in the mouth and bill. The
majority of caches by pinyon and scrub
jays are singleseeded, and are located
in the transition zone between mineral
soil and the overlying organic material.
Although rodents are known to cache
seed, they should not be considered
effective seed dispersers because caches
are located in middens or underground
chambers where conditions are not
suitable for germination or seedling
establishment. Instead, rodents, such as
cliff chipmunks, pinyon mice, and
woodrats, are major predators, caching
as much as 35 to 70 liters (1 to 2 bu)
of good seed. Furthermore, limited
data indicate that rodents consume large
quantities of seeds taken from bird
caches.
It has been suggested that pinyon trees
and seed eating birds have evolved
coadaptive traits that enhance survival
of both organisms. The seed dispersing
and caching behavior of birds appears
related to certain traits of the trees:
large, thin-coated seeds with high
energy values, different colored
seedcoats that aid visually oriented
seed harvesters to distinguish edible
from aborted seeds, upward orientation
of cone and scale angle for increased
seed visibility, and prolonged seed
retention in open cones because of cone
orientation and deep depressions and
small flanges on cone scales.
Furthermore, the mutual dependence of
birds and trees appears more complex
than just their respective roles of seed
dispersing and food providing agents.
Gonadal activity of pinyon jays, for
example, is increased before the
breeding season by the combined effect
of photoperiod, the appearance of cones,
and a diet of seeds. (1) |