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1995 ; Vesk & Westoby 2004 ; Falster & Westoby 2005 ; Knox & Clarke 2005 ).
The extent to which these differences affect competitive interactions between
obligate seeders and facultative seeders is a matter of some controversy (Keith &
Myerscough 1993 ; Clarke et al. 2005 ; Wardell-Johnson et al. 2007 ) and is discussed
further in Chapter 9 .
Seed Supply Constraints
The bulk of species have persistent seedbanks in which there is some seed carry-
over from year to year and seeds are usually available for germination in situ
following fire (Bell 1999 ; Auld et al. 2000 ; Pausas et al. 2004b ; Enright et al . 2007 ;
Lamont et al. 2007 ; Pausas & Bradstock 2007 ). The formation of persistent in situ
seedbanks is tied to deep dormancy and relatively short range dispersal for
the bulk of the seed pool (long-distance dispersal of some of these is known,
Bradstock et al. 1996 ; Lamont et al. 2007 ). Such species are not reliant on annually
seeking gaps for establishment, but instead are dependent on disturbance in situ
for the provision of gaps.
Seedbanks stored in the canopy through serotiny (see Fig. 3.5 and Table 9.2 ), a
form of bradyspory, represent about 25% of the postfire seeders in MTC ecosys-
tems in Australia (Lamont et al. 1991 , 2007 ). Soil-stored seedbanks comprise
about 70% (Keith et al. 2002 ; Pausas et al. 2004b ). Soil seedbanks of woody
species may accumulate more rapidly and have greater longevity than their
serotinous counterparts. This is due to several factors. Species with soil seed
storage tend to mature earlier (Auld 1987 ; Bradstock & O'Connell 1988 ; Keith
et al. 2002 ; Abbott & Burrows 2003 ; Burrows et al. 2008 ). Also, soil-stored
seedbanks possess dormancy mechanisms that may enable seeds to outlive parent
plants (Bell 1999 ; Auld et al. 2000 ; Whelan et al. 2002 ; Wills & Read 2007 ). In
addition, there is some evidence that soil seedbanks are not entirely exhausted by
postfire germination and may persist through repeated fires (Whelan et al. 2002 ;
Tozer & Auld 2006 ); see also Chapter 7 .
Canopy storage in obligate seeder species of shrubs generally take 4-10 yrs to be
initiated. Slower rates of maturation occur in extreme habitats such as rock
outcrops (Clarke 2002a ; Yates et al. 2003a , 2003b , 2007 ; Burrows et al. 2008 )
and in gymnosperms such as Callitris spp. (Bradstock & Cohn 2002b ). Canopy
storage in facultative resprouters is replenished more rapidly because flowering
begins very early in resprouts and thus this secondary juvenile period is much
shorter than the primary juvenile period following germination (Bradstock 1990 ;
Lamont et al. 1998 , 2007 ). Compared with congeneric obligate seeders, seed crops
of facultative resprouters with canopy storage also tend to be lower (but not
always, see Enright et al. 2007 )(Zammit&Westoby 1988 ;Bell 2001 ; Pausas et al.
2004b ;Lamont et al. 2007 ). Serotinous storage generally reaches maximum levels
>
10 yrs postfire, with rate of growth of storage declining at this time, due to losses
from predation (vertebrate and invertebrate) and spontaneous release (Gill &
McMahon 1986 ; Bradstock & O'Connell 1988 ;Bradstock 1990 ;Lamont&
Groom 1998 ; Lamont et al. 2007 ).
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