Rh. Okland, POPULATION BIOLOGY OF THE CLONAL MOSS HYLOCOMIUM SPLENDENS IN NORWEGIAN BOREAL SPRUCE FORESTS .1. DEMOGRAPHY, Journal of Ecology, 83(4), 1995, pp. 697-712
1 Hylocomium splendens is a perennial clonal moss with annual periodic
ity in the emergence of new modules. New growing points usually emerge
in spring and develop into mature segments their second summer. Popul
ation increase is brought about by ramification (branching) of the you
ngest segment or by regeneration from older parts of the shoot system.
2 A technique for nondestructive re-identification of segments in per
manent plots is devised. Precise demographic information on 9206 growi
ng points and 7602 mature segments was obtained by following 119 local
populations in seven Norwegian reference areas for monitoring during
3 years. The dry weight of each mature segment was estimated from morp
hological field measurements. 3 The potential of each mature segment f
or production of new growing points was strongly size-dependent; c. 40
% of the variation in a segment's size was explained by the size of it
s mother segment. Integration within segment chains was largely restri
cted to neighbouring segments. The number offspring growing points (by
ramification as well as regeneration) was positively size-dependent,
while termination risk was negatively size-dependent. Mature segments
produced by regeneration were smaller, while segments produced by rami
fication were larger than the average segment. Strong apical dominance
was demonstrated. 4 Mortality and branching rates, averaged over popu
lations, areas and years, are reported. Annual recruitment was 32.3%;
23.3% by regeneration, 7.5% by ramification, and 1.5% by accidentally
introduced shoots. Annual mortality was 19.7%. 6.8% by loss, 1.9% by g
razing, and 11.0% by termination for other reasons.5 Demographic data
for growing points and mature segments were analysed separately by sta
ge-structured transition matrix models. The total population increased
every year (average lambda = 1.10-1.11). Vital rates were strongly si
ze-dependent. The contributions of ramification and regeneration to in
crease in lambda are equal as ramification occurred in large segments
with high probability for survival. 6 Ramification is considered impor
tant for maintenance of size and risk-spreading among ramets under fav
ourable conditions, while high regenerative ability facilitates resili
ence to, and recovery after, disturbance. The ecological significance
of size is discussed. 7 Hylocomium splendens is closely similar to clo
nal forest vascular plants in major demographic traits, e.g. size-depe
ndence of fitness and branching patterns.