Margalef's
mandala maps phy
toplan
kton species in
to a phase space defined by
turbulence (
A) and nu
trien
t concen
tra
tions (
Ni);
these are
the hard axes. The permu
ta
tions of high and low
A and high and low
Ni divide
the space in
to four domains. Sof
t axes indica
te some ecological dynamics. A main sequence shows
the normal course of phy
toplan
kton succession;
the
r-K axis of MacAr
thur and Wilson runs parallel
to i
t. An al
terna
tive successional sequence leads
to
the low
A-high
Ni domain in
to which many red
tide species are mapped. As
tronomical and biological
time are implici
t. A ma
thema
tical
transforma
tion of
the
mandala (ro
ta
tion) lin
ks i
t to
the classical bloom models of Sverdrup (
time) and Kiers
tead and Slobod
kin (space).
Both rarity and the propensity to form red tides are considered to be species characters, meaning that maximum population abundance can be a target of natural selection. Equally, both the unpredictable appearance of bloom species and their short-lived appearances may be species characters. There may be a correlation too between these features and long-lived dormant stages in the life-cycle; then the vegetative planktonic phase is the 鈥榳eak link鈥?in the life-cycle. Red tides are thus due to species which have evolved suites of traits which result in specific demographic strategies.