13Amaro Forests - Chap 11 1/8/03 11:52 am Page 129
129 Temporal Dependence of Pollen Cone Production
observed and expected frequencies is quite reasonable, yet a Kolmogorov–
Smirnov test (Conover, 1980) indicated statistical significance of their differ-
ˆ
ences (maximum absolute difference D = 0.05, P
< 0.01). The observed
max
‘surplus’ of trees with pollen cones in 8 of the years accounted for most of the
discrepancies.
Discussion and Conclusions
Statistical tests for the order of a finite binary Markov chain are most powerful when
the frequencies of zeroes and ones are in the intermediate range between 0.3 and 0.6
(Billingsley, 1961). When jack pine carries pollen cones, on average of three out of
every 4 years, the number of non-estimable or poorly estimated transition probabili-
ties becomes a non-trivial issue. Also, the 9-year records used here are probably at
the lower limit for statistical inference.
Our results do not clearly reject the null hypothesis of temporal independence
of pollen cone pr
oduction, at least not at the individual tree level. Conditional on
the location effect, it appears reasonable to accept, as a working model, the notion of
temporal independence. A first-order model, however, would, from an ecological
viewpoint, be more realistic. Annual variations in growing conditions and distur-
bances of the physiological conditions can be expected to influence the tree beyond
the year in which they occur. A precocious pollen cone production amounts to a sig-
nificant carbon sink for the tree. A feedback control of pollen cone production gov-
erned by environmental cues would seem a reasonable hypothesis. Periodicity of
seed set is the norm (Greene and Johnson, 1994); production of pollen cones is
expected to show a similar pattern.
Although we settled for a binomial model to explain the observed tr
ee level
frequencies of years with pollen cones, we recognize that a hierarchical model, with
a first-order Markov chain at the first level, a Dirichlet distribution describing the
mixing distribution of the transition probabilities within a location (p
00
, p
01
, p
10
, p
11
)
at the second level, and at the third level a multivariate distribution of the
Dirichlet distribution parameters to capture the effects of locations, might provide a
more satisfactory fit overall. A practical reason for assuming order 1 is that order 0
becomes a special case, which could be tested under a hierarchical model, in much
the same way as done here. We are currently exploring hierarchical models.
References
Benzie, J.W. (1977) Manager’s Handbook for Jack Pine in the Northern Central States. NC-32, USDA
Forest Service.
Billingsley, P. (1961) Statistical Inference for Markov Processes. University of Chicago Press,
Chicago, Illinois, 194 pp.
Collett, D. (1991) Modelling Binary Data. Chapman and Hall, London, 369 pp.
Conover, W.J. (1980) Practical Nonparametric Statistics. Wiley, New York, 493 pp.
Conway, B.E., Leefers, L.A. and McCullough, D.G. (1999) Yield and financial losses associated
with a jack pine budworm outbreak in Michigan and the implications for management.
Canadian Journal of Forest Research 29, 382–392.
Greene, D.F. and Johnson, A.E. (1994) Estimating the mean annual seed production of trees.
Ecology 77, 642–647.
Griffiths, D.A. (1973) Maximum likelihood estimation for the beta-binomial distribution and an
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