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of Lonicera , Clematis , Smilax and Rubia are also common, although these taxa
occur in evergreen oak woodlands as well.
These maquis communities are almost exclusively dominated by postfire obli-
gate resprouters and the proportion of woody species that are killed and fail to
resprout after fire is generally very low. As a consequence this vegetation regener-
ates rapidly after fire. Most of these shrubs have vertebrate-dispersed fruits, often
fleshy ( Table 4.3 ), and consequently they are dispersed relatively long distances
(Jordano et al. 2007 ; Pons & Pausas 2007 ). These obligate resprouting species
avoid the stressful summer drought by means of deep roots that gain access to
water through soil cracks. Thus, many of the seedlings are dependent on establish-
ment in favorable mesic microsites (see Chapter 9 ) and they often make an early
investment in roots (Lloret et al. 1999b ; Paula & Pausas 2011 ).
Perhaps one of the most typical broadleaf shrubs in the Mediterranean Basin is
Quercus coccifera , known as Kermes oak, which is an evergreen species that
resprouts vigorously from rhizomes after fire (Malanson & Trabaud 1988 ). For-
mations dominated by this shrub are locally termed garrigue ( Box 4.1 ). Although
this species is resilient to frequent fire, there is a reduction in productivity
when subjected to very short fire intervals (Delitti et al. 2005 ). The floristic compos-
ition of these communities also changes very little with different fire regimes
(Trabaud & Lepart 1980 , 1981 ). In the western part of the basin, Q. coccifera tends
to be a multistemmed shrub, often forming large and imbricate carpets, but occa-
sionally forming low woodlands on moister sites, for example in North Africa
(Charco 1999 ). In the eastern part of the basin, Q. coccifera is very often a woodland
tree and this form is often considered a distinct species, Q. calliprinos .
A few widespread species in these shrublands have highly reduced leaves
and photosynthetic stems, sometimes flattened to form phylloclades. These
include species of Ruscus (with broad sclerophyllous phylloclades) and species of
Asparagus (with thin, sometimes thorny, phylloclades). These taxa have fleshy
fruits and are obligate resprouters after fire. The native dwarf palm Chamaerops
humilis may also appear in these western shrublands; its leaves protect the apical
meristem from fire and thus it survives crown fires. Indeed, it may become
dominant in zones with very high fire recurrence, and also under very high grazing
pressure. The only other palm native to the region, Phoenix theophrasti , also
survives fire but it occurs on very few sites in the eastern part of the basin (Boydak
1985 ; Barrow 1998 ).
Intermixed with the broadleaf shrubs, some small-leaved species may also
occur, for example mostly resprouting Erica species or Juniperus oxycedrus , which
has both resprouting and non-resprouting populations (Pausas et al. 2008 ); the
non-resprouting Juniperus phoenicea appears only on sites that have had long fire-
free periods or that are fire protected, such as rocky outcrops and cliffs. Other
small-leaved species that may appear together with broadleaf shrubs, especially on
open microsites, are species of Lamiaceae, Cistaceae and Fabaceae. Many of these
have the capacity to recruit seedlings after fire but are much more dominant in
other shrubland formations.
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