Plant Ecology (2005) 181:127–137
Springer 2005

DOI 10.1007/s11258-005-5321-x
0

Gypsum physical soil crusts and the existence of gypsophytes in semi-arid

central Spain

Roberto L. Romão1 and Adrián Escudero2,*

1
Universidade Estadual de Feira de Santana. Laboratório de Ecologia Evolutiva da, Av. Universitária, s/n.
44031-460 Feira de Santana – BA, Brasil; 2Biodiversity and Conservation Group. E.S.C.E.T. Universidad
Rey Juan Carlos. C/. Tulipán s/n. Móstoles E-28933. Spain; *Author for correspondence (e-mail: adrian.
escudero@urjc.es; phone: +34-91-4887070)

Received 22 January 2004; accepted in revised form 7 April 2005

Key words: Edaphic specialist, Emergence, Physical soil crusts, Seedling growth, Seedling survival

Abstract

Probably gypsophytes are the most interesting set of edaphic specialists of arid and semiarid climates.
Despite they conform a global biodiversity priority, there are almost no information about those adaptive
traits that confer such a specialised behaviour. Our broad hypothesis is that gypsophytes are ‘‘refuge-
endemics’’ that are able to grow on gypsum soils due to their ability to surpass extremely hard gypsum soil
physical crust during emergence. With this in mind we have conducted an experimental approach com-
bining field and greenhouse assays. Seeds from two gypsophytes, genuine and widely distributed in the
Iberian Peninsula gypsophytes (Helianthemum squamatum and Lepidium subulatum) and one gypsovag
(Teucrium capitatum), a generalist plant that can also grow on gypsum soils were used in our experiments.
Two complementary experimental approaches were conducted. The first involved a field experiment in
which the presence or absence of the physical crust together with the sowing date were manipulated and a
greenhouse experiment in which the irrigation amount and the types of soil were controlled. Variables of
interest were the percentage of germination, growth and survival. In the field experiment we found a
significant decrease in the final germination of the gypsovag in the plots with intact crusts. On the other
hand, H. squamatum is able to grow in the three tested soils, despite higher survival and growth on genuine
gypsum soils. Our results confirm the hypothesis that gypsum edaphic specialists base their behaviour to a
great extent on the ability to surpass extremely hard gypsum surface crusts, although this seems a marginal
adaptive trait as shown by the capability to grow on a complete array of soils and the negative effect of the
crust along the earlier development life stages of gypsophytes. Furthermore, a gypsovag such as Teucrium
capitatum presents extreme difficulties to emerge on non-disturbed gypsum physical crusts but once sur-
passed its growth and survival is not limited.

Introduction Garcı́a-Moya 1989; Mota et al. 2003). These

Gypsophytes probably represent the most con-
spicuous and diversified set of arid and semiarid
endemic plants (Johnston 1941; Parsons 1976;
Powell and Turner 1977; Meyer 1986; Meyer and
plants exclusively grow in gypsum (hydrous cal-
cium sulfate) soils under arid or semiarid climates.
Surprisingly, such type of specialists have received
remarkably little study (Meyer 1986; Meyer et al.
1992; Escudero et al. 1999, 2000) in spite of
128
gypsisols which are characterised by a gypsum
content over 5%, extending over one million km2
in the world (Verheye and Boyadgiev 1997). In
addition to this, most of these plants conforms a
global biodiversity priority because most of them
are narrowly distributed and seriously threatened
(Meyer 1986). In these sense, Mediterranean gyp-
sum ecosystems are considered one of the most
threatened habitats in the Mediterranean Basin
(Gómez-Campo 1987; European Community
1992; Mota et al. 2003).
Historically, gypsophily has been related to the
ability of gypsophytes to persist under certain
chemical restrictions of gypsum soils (Duvigneaud
and Denaeyer-De Smet 1966, 1968; Boukhris and
Loissant 1970; Cannon 1971). However several
field studies have shown little evidences of this type
of restriction (Meyer 1986; Meyer and Garcı́a
Moya 1989; Meyer et al. 1992; Verheye and Boy-
adgiev 1997). It has been also suggested that the
specialist behaviour of gypsophytes lies to a great
extent in their capacity to surpass extremely hard
gypsum physical surface crusts immediately after
germination (Campbell and Campbell 1938; Meyer
1986; Escudero et al. 1999, 2000). So, emerging
gypsophytes surpass the gypsum soil crust,
whereas non-gysophytes are unable to. Our spe-
cific hypothesis is that gypsophytes take selective
advantage due to this capability, whereas gypso-
vags are plants that able to grow on gypsum but
not exclusively, only recruit individuals in the
vicinity of standing gypsophytes where gypsum
physical crusts are narrower; probably via facili-
tation (Escudero et al. 2000). It is widely known
that gypsum crusts are notoriously thicker and
harder than those found in other arid and semiarid
habitats (Verherye and Boyadgiev 1997). Ad-
ditionally, during the past few years we have
accumulated field evidences of notorious gypso-
phyte emergences and survival on extremely well
conserved hard-crusted gypsum surfaces (Escu-
dero et al. 1999; 2000; Romão 2003).
Our broad hypotheses are (1) the existence of a
profound link between gypsophily and the pres-
ence of thick non-perturbed gypsum physical
crusts that only can be surpassed by genuine gy-
psophytes at least on massive gypsum outcrops,
and (2) the fact this behaviour is a marginal
adaptive trait, so gypsophytes constitute ‘‘refuge’’
endemics (in the sense of Gankin and Major 1964)
and consequently are able to grow on other soils
where they are competitively excluded under
natural conditions. In order to test these two
complementary hypotheses, we have conducted a
two-fold experimental approach combining field
and greenhouse experiments. If our gypsum
physical crust hypothesis is correct, gypsophytes
should germinate, grow and survive on crusted
soils whereas gypsovags, plants that marginally
can grow on gypsum soils, should perform much
better on finely raked gypsum soils (crust experi-
mentally crushed). On the other hand, if gypso-
phytes are not soil specialists and exploit gypsum
environments as marginal habitats where they are
competitive, they should germinate and recruit
individuals not only on gypsum soils but also on
other types of soil.
Methods
We have selected three conspicuous shrubby plants
of the gypsum communities of central Spain (Rivas-
Martı́nez and Costa 1970; Rubio et al. 1994). He-
lianthemum squamatum (L.) Cours (Cistaceae) and
Lepidium subulatum L. (Cruciferae) are small
shrubs (20 – 60 cm) that exclusively grow on gyp-
sum outcrops. These two plants are typical ele-
ments of the so-called Gypsophiletalia sthrutii
communities, which are endemic but widely dis-
tributed on gypsum outcrops under semiarid con-
ditions in the Iberian Peninsula (Loidi and
Fernández-González 1994). However, Teucrium
capitatum L. (Labiatae) which also is a small shrub
can grow on gypsum soils but preferentially on the
widely distributed calcareous soils from the Eastern
half of the Iberian Peninsula. This gypsovag has
been selected because it is one of the most out-
standing elements both in gypsum and calcareous
soils in the vicinity of our field experimental site.
Experimental approach
Experiment one – The field study site was located
in Chinchón, Madrid province, within an excep-
tionally well-conserved remnant of a southern
exposed gypsum habitat (4011¢ N, 335¢ W,
550 m a.s.l.). The climate is upper semiarid with an
annual average rainfall of 415 mm but almost no
rainfall in summer. Mean daily maximum and
minimum temperatures in January are 9.6 C and

0.6 C, respectively, and 32.7 C and 9.6 C in July.
Soils are Calcic gypsisols developed over gypsum
parental rocks (Monturiol and Alcalá del Olmo
1990) with gypsum contents above 50%. Vegeta-
tion is conformed by conspicuous vegetated pat-
ches dominated by creeping chamaephytes
interspersed on skeleton crusted surfaces covered
by specialised lichens. Cover of perennial plants is
20% in the 1000 m2 where the experimental plots
were set. The dwarf scrub community has been
named Herniario fruticosae-Teucrietum pumili
(Rivas-Martı́nez and Costa 1970). The bare zone is
covered by a biological crust (50%) and some
gypsum crystals (5%). Soil biological crust is
dominated by Diploschistes diacapsis, Squamarina
cartilaginea together with some gypsum specialised
lichens such as Toninia sedifolia, Fulgensia subb-
racteata and Psora decipiens and with almost non
fix-nitrogen elements such as Collema crispum.
This biological crust is extremely hard with an
average strength in May of 1230 kPa (SE = 350,
n = 20) measured with a portable penetromer
(RIMIK CP-20, Australia). During the summer
drought the physical crust reaches significantly
higher values (Escudero, personal observation).
Experimental plots consist 30 cm · 30 cm qua-
drates randomly distributed on bare (without
perennial cover) and flat (<5%) soil surfaces.
Plots were randomly assigned to two contrasting
and fixed treatments. In the first, named ‘‘intact
crust’’, the soil surface was unaltered, whereas in
the ‘‘disturbed crust’’, soil surface was evenly
raked to destroy the gypsum crust in January
2000. Finely divided microphytic rests were not
retired from the quadrates. Six plots per treatment
and plant species were considered. Twenty five
sowing circles (15 mm in diameter) were regularly
distributed in a transparent plastic sheet in order
to conform a 5 cm side-cell grid. The plastic sheet
was used for sowing and monitoring purposes but
not left on soil. Three seeds per cell grid per species
were sown in middle January within each plastic
hole conforming a set of 75 seeds per quadrate.
Seeds were separated to avoid interferences during
the emergence. Seeds were sown on the soil surface
and not buried. Previous works on gypsophyte
germination responses showed that final germina-
tion percentage reached more than 50% in the
three considered species (Escudero et al. 1997), so
the number of sown seeds should guarantee at
least one seedling per sowing point once emerged.
129
Sowing was conducted in January immediately
before field emergences under natural conditions
(Escudero et al. 1997). Seeds were collected during
the previous summer from 20 to 30 plants per
species randomly selected within the experimental
area and stored at 6 C in darkness in hermetically
sealed containers (see Escudero et al. 1997). Before
sowing, seeds were soaked in distilled water in
order to promote the development of seed muci-
lage. The existence of mucilaginous seed coats is a
widely extended trait in gypsum environments and
these three species have mucilaginous coats
(Escudero et al. 1997). Such a type of seed trait
determinates both a favourable environment for
germination and an efficient mechanism to avoid
washing, so seeds remain anchored at the sowing
point. In the case of the ‘intact crust’ plots lichens
were carefully eliminated in the sowing hole when
necessary to avoid bias due to the nature of the
lichen species.
Field germination was regularly recorded till
October because the most critical period for
seedling survival corresponds to the first summer
drought. Second and third germination per hole
were noted but seedlings were immediately clipped
to avoid pernicious seedling competition. Seedling
survival and growth was also recorded and mea-
sured as the number of leaves (Helianthemum
squamatum) and total length (Lepidum subulatum
and Teucrium pumilum) at each census.
As shown in (Escudero et al. 1999) survival is
higher for the first Helianthemum squamatum
cohort. In order to evaluate the relationships
between germination time and crust constraints we
conducted a similar experiment with Helianthe-
mum squamatum but delaying the sowing time to
the first week of March. The new source of vari-
ation in the corresponding linear models was the
emerging time (two fixed levels: January and
March sowing). During the germination window
(January to May) the strength of the gypsum crust
remains rather constant and relatively low.
Experiment two – In a parallel greenhouse
experiment the specialist behaviour of a widely
distributed gypsophyte such as Helianthemum
squamatum was also surveyed. A two-way factorial
experiment was conducted. Soil type was consid-
ered the first source of variation with three fixed
levels, gypsum soils and calcareous soils which
were collected in the field in the vicinity of the
above mentioned locality (soil data in Romão
130
2003), and an enriched commercial substratum
(COMPO SANA Universal). The second source of
variation was the amount of irrigated water. It
must be reminded that the gypsophile behaviour is
related to the amount of rainfall, being confined to
arid and semiarid climate conditions (Rivas-
Martı́nez and Costa 1970). Two fixed levels were
considered, a control which mimics the amount of
rainfall during spring and early summer in our
field locality (12 ml per pot) and a second treat-
ment which double this last amount (one irrigation
per week). Twenty-five pots of 10 · 10 · 10 cm
were filled per treatment combination. Five seeds
were sown per pot in January. This density is
similar to that found under natural conditions in
favourable microsites (Escudero et al. 1999).
Provenance and conservation of seeds were similar
to the above mentioned experiment. Germination,
survival and growth were also weekly recorded
during 6 months.
Statistical analysis
Experiment one – Normality of the variables –
percentage of germination, survival at June and
survival at September per plot was studied
through the Kolgomorov – Smirnov test
(p < 0.1). Appropriate transformations were
applied to improve normality when necessary.
Linear models were independently conducted for
each species with two fixed treatment, crust (1
d.f.) and germination date (1 d.f.). When nor-
mality was not reached, non-parametric tests
were conducted.
Experiment two – Normality of variables were
also checked – average plant size per pot in June,
size in September, average number of leaves per
plant and pot in June and number of leaves per
plant in June – and two-way anovas was con-
ducted on log-transformed data. To account
statistically for possible differences among pots in
their response to experimental treatments due to
competition, the number of seedlings per pot was
included in the model as a covariate (substra-
tum*water*number of seedlings per pot). As this
study focuses on the consequences of experimental
manipulations, significance tests associated with
covariate in models are not considered.
The significance of the effects of sowing sub-
stratum (substratum) and irrigation (water) on the
probability of one seed yielding an emerged seed-
ling per pot (‘‘seedling emergence rate’’), proba-
bility of survival at June and probability of
survival at September were also assessed by fitting
generalised linear models. These variables were
modelled as binomial response due to their nature
and difficulties to normalise, using logits as the
link function. As variables were sequentially in-
cluded, we tested different sequences and nested
models (Mc Cullagh and Nelder 1989) and sub-
sequently compared. Models were adjusted using
S-Plus 2000.
Results
Experiment one
As shown in Figure 1 and Table 1, Lepidium
subulatum germination did not show differences
between treatments (average final germination =
43.38% ± 2.82), nor survival was affected (sur-
vival in May = 72.39% ± 8.09; in June =
35.31% ± 8.17; and in September = 7.17% ±
4.35). On the other hand Teucrium capitatum seeds
emerged significantly better without crust (final
germination = 29.11% ± 4.11) than on crusted
soils (5.11% ± 2.31) (Figure 1 and Table 1).
However survival was rather similar between
treatments (survival in May = 68.85% ± 7.37; in
June = 48.40% ± 8.84, and in September =
8.98% ± 3.16). Germination of Helianthemum
squamatum seeds only showed significant differ-
ences for the sowing treatment (Figure 1 and
Table 2), being higher the final germination for the
first cohort (final germination = 53.00% ± 3.29
for the January cohort and 33.77% ± 2.37 for the
March cohort). After germination, survival did
not present significant differences neither for the
crusted treatment nor for the sowing moment
(survival in May = 71.14% ± 3.64; in
June = 54.73% ± 3.96; and in September =
16.06% ± 3.22).
Size in June was extremely different in the case
of L. subulatum (U = 1633, p < 0.0001) being
higher in the disturbed and crushed crust treat-
ment, marginally different in H. squamatum
(U = 1089, p = 0.036) following a similar trend,
and rather similar in the case of T. capitatum
(U = 145, p = 0.487)
 131

Figure 1. Mean value and standard error for the original data of germination in disturbed and intact crusts and taking into account the
sowing date for H. squamatum (field experiment).

Table 1. Results from the experiment I (field conditions) for Lepidium subulatum (gypsophyte) and Teucrium capitatum (gypsovag).

Plant Species Variable df F p-value

Lepidium subulatum Germination percentage 1,10 0.685 0.427

May Survival 1,10 0.268 0.618
June Survival 1,10 3.09 0.108
September Survival v 2 = 3.578 0.058
Teucrium capitatum Germination percentage 1,10 13.5 0.000
May Survival 1,10 2.10 0.177
June Survival 1,10 2.06 0.181
September Survival v 2 = 0.117 0.731

Survival in September could not be properly transformed, so a non-parametric test was conducted.
132

Table 2. Results for H.squamatum in the experiment I (field conditions). Models included crust treatment (intact and disturbed crust),
and sowing cohort (January and March).

Variable Source of variation df F p-value

Germination percentage Crust treatment 1, 20 1.12 0.301

Sowing cohort 1, 20 14.62 0.000
Interaction 1, 20 3.06 0.095
May Survival Crust treatment 1, 20 3.76 0.066
Sowing cohort 1, 20 0.61 0.000
Interaction 1, 20 1.69 0.207
June Survival Crust treatment 1, 20 0.000 0.992
Sowing cohort 1, 20 0.865 0.363
Interaction 1, 20 1.007 0.327
September Survival Crust treatment 1, 20 0.012 0.914
Sowing cohort 1, 20 2.275 0.147
Interaction 1, 20 0.593 0.450

Experiment two 1.00 ± 0.40; log transformed), but marginally

Treatments showed no differences in the Helian-
themum squamatum average size per pot in June
(average size per pot – log transformed = 0.71 ±
0.17) and only for substratum in size in September
(Table 3). Pairwise comparisons between substra-
tum levels clearly show that size in enriched com-
mercial soils becomes significantly larger
(1.57 ± 0.64; log-transformed p < 0.000) than in
gypsum soils (log-transformed size = 0.88 ± 0.17)
or calcareous soils (0.87 ± 0.15). These last two
levels do not exert significant differences. We have
also no detected differences in the average number
of leaves per plant and pot in June (mean =
higher in the enriched soil also in September (Ta-
ble 3).
Maximum likelihood analysis of variance of
dependent variables for the effects of sowing sub-
stratum (Substratum) and irrigation (Water) were
obtained adjusting generalised models as binomial
responses (Table 4). Seedling emergence rate
showed no differences among treatments. How-
ever, survival in June and survival in September
showed highly significant differences among sub-
strata (Figure 2). Survival in gypsum soils is
always significantly higher, mainly in relation to
enriched soils. A significant interaction between
irrigation and substratum was detected in both

Table 3. Results of the experiment II (greenhouse conditions).

Variable Source of variation df F p-value

Size in June Substratum 2,181 0.424 0.655

Water 1,181 0.525 0.470
Substratum*Water 2,181 2.117 0.124
Size in September Substratum 2,181 4.54 0.012
Water 1,181 0.616 0.433
Substratum*Water 2,181 0.397 0.673
Leaves in June Substratum 2,181 0.092 0.912
Water 1,181 1.623 0.203
Substratum*Water 2,181 2.363 0.097
Leaves in September Substratum 2,181 4.011 0.020
Water 1,181 4.392 0.047
Substratum*Water 2,181 2.560 0.080

The treatments included were the different soil substrata (gypsum, calcareous soils and enriched commercial substratum) and water
(two levels of irrigation). All variables were log transformed to fulfil normality criteria. Average values per pot (n = 50 per treatment
combination).

Table 4. Maximum likelihood analysis of variance of dependent variables from experiment II for the effects of sowing substratum
(Substratum) and irrigation (Water) were obtained adjusting generalised models as binomial responses (logit as link function).
Variable Source of variation
Seedling emergence rate Substratum
Water
Substratum*Water
Survival at June Substratum
Water
Substratum*Water
Survival at September Substratum
Water
Substratum*Water
cases, mainly because under double irrigation
treatment, survival in calcareous soils is signifi-
cantly lower (Figure 2). In the case of the survival
in September the reduced model (without the
interaction) was not a good reduction of the sat-
urated model (v2=15.07, p = 0.0005).
Discussion
Many authors have highlighted the relevance of
biological crusts in arid systems (Watson 1979;
Belnap et al. 2001). They redistribute water (Eld-
ridge et al. 2000; Maestre et al. 2002), enhance soil
stability (Belnap and Gillette 1998) and the activity
of microphytes improves soil resource levels (Bel-
nap and Harper 1995; Zaady et al. 1998) deter-
mining ecosystem function and dynamics (West
1990), and resilience, facilitating ecosystem recov-
ery after severe droughts (Aranibar et al. 2003).
However several sowing experiments have shown
an important reduction in germination on undis-
turbed crusts (Zaady et al. 1997). It is also known
that the development of desert physical crusts is one
of the most limiting factors for the emergence of
new seedlings under field conditions (Hanks and
Thorp 1956; Harper et al. 1965; Cary and Evans
1974). Additionally, Sylla (1987) and Johansen
(1993) have suggested that the inhibition mecha-
nism in microphytic crust is related to the difficul-
ties of the penetration of radicles and seeds into the
soil below the crust layer. Prasse and Bornkann
(2000) pointed out that the effect of crusts on plant
is extremely diverse and probably include a trade-
off between negative effects at the germination stage
and the above mentioned benefitial features. Our
results clearly support the physical hypothesis
133
df Chi-square p-value
2 1.269 0.530
1 0.227 0.633
2 1.59 0.450
2 42.13 0.0000
1 0.032 0.856
2 15.07 0.0005
2 39.29 0.0000
1 1.29 0.254
2 8.58 0.013
about the origin of gypsophily (see Meyer 1986;
Escudero et al. 1999). Field evidences had previ-
ously shown that the two studied gypsophytes,
Lepidium subulatum and Helianthemum squamatum
are able to recruit in extremely hard gypsum sur-
faces (>1500 kPa in gypsum crests) under a wide
array of climatic conditions (Escudero et al. 1999;
2000). Such observational data seems to support
for our experimental results. They confirm that
emerging radicles of gypsophytes can surpass the
gypsum physical crust without significant difficul-
ties. However, gypsovag Teucrium capitatum has
enormous difficulties to emerge (only 5% in aver-
age) under crusted conditions whereas not on un-
crusted soils. Although Bridges and Burnham
(1980) pointed out that roots of generalists plants
were not able to penetrate in soils with more than
25% of gypsum (gypsisols) because the soil mate-
rials lack plasticity, cohesion and aggregation, and
is completely mechanically unstable, we consider
that the most relevant fact is the inability of
emerging seedlings to surpass the gypsum crusts
and not the lately seedling or developed plant. It is
necessary to note that the tested effect is only re-
lated to the extremely hard physical constraint of
the gypsum crust, because the filtering effect of the
biological components of the crust on any life stage
of vascular plants remain unknown. In any case this
non-explored effect should be additive because well
conserved biological crusts always present extreme
strength values. Lichen crust cover appears high
and positively correlated with the strength of the
gypsum crust (Tarazona et al. 1980).
Other two evidences reinforce the physical
hypothesis: first, Merlo et al. (1997) and Romão
(2003) have not found significant differences in
laboratory germination under a complete set of
134
Figure 2. Mean value and standard error for the original data (not our binomial model) of survival in June and September in
experiment two (greenhouse experiment).
gypsum dilutions, showing that the presence of
gypsum and or calcium content do not alter the
germinability in these two gypsophytes, and sec-
ond, gypsophytes are confined to arid or semi-arid
climates where gypsum crusts exclusively appear
(Rivas-Martı́nez and Costa 1970; Meyer 1986).
Under more mesic climates, gypsum physical
crusts do not appear and the gypsum outcrops,
sometimes with percentages of sulfates in soil far
above 50%, are covered with calcicole vegetation
(Rivas-Martı́nez and Costa 1970). On the other
hand, Teucrium capitatumis relatively abundant in
the gypsum outcrops of the studied area, but
confined to vegetated patches where genuine
gypsophytes are dominant (see Romão 2003).
There are no multi-species patches dominated by
T. capitatum nor isolated individuals in non-dis-
turbed community (Romão 2003).
As previously reported (Escudero et al. 1999),
timing of emergence is a key factor for growth and
consequently for survival in many desert plants
(see Fowler 1988; Eldridge and Westoby 1991), but

as shown here, also for the germination of
H. squamatum. Germination is significantly higher
for the first sowing cohort. Factors underlining
this behaviour might be related to a temperature
control because most of the Iberian gypsophytes
present optim temperature for germination rang-
ing between 5 C and 15 C (Escudero et al. 1997)
and soil’s water contents remain rather similar
along all the autumn-early spring period. This
widely extended trait in Mediterranean plants
(Thanos et al. 1992; Bell et al. 1993) must guar-
antee an optimum seedling survival.
On the other hand, no significant differences in
survival (in May, in June and even in September)
appear in any of the three studied plant species,
two genuine gypsophytes and a gypsovag. This
fact suggests that once the crust is surpassed the
chance for survival is maintained both for gypso-
phytes and gypsovags in the two experimental
scenarios. As expected the most limiting factor for
survival is the summer drought as previously
reported under non-experimental field conditions
(Escudero et al. 1999; 2000), even for other Iberian
gypsophytes such as Gypsophila struthium subsp.
struthium (Solá, personal communication).
Although not included, logistic models for survival
in July (mortality peak) for the three studied spe-
cies were highly significant and included the seed-
ling size as predictor with positive coefficients (see
also Escudero et al. 2000). Anyway, seedlings
better grow on disturbed crust in the case of L.
subulatum but also marginally in H. squamatum.
This fact points out the existence of a negative
effect of physical gypsum crust during the earlier
growth stages on both gypsophytes, suggesting
that the population mediator role of the physical
gypsum crust is not only confined to emergence
(see Bridges and Burnham 1980).
H. squamatum seems to conform a ‘‘refuge-
model’’ endemic (sensu Gankin and Major 1964;
Proctor and Woodell 1975) because seeds of
H.squamatum germinate without significant dif-
ferences in the three tested soils and can efficiently
grow on non-gypsum soils. This result would be in
agreement with Boukhris and Loissant (1970),
who pointed out the capability of gypsophytes to
grow on several types of soil. However, we have
found significant differences in survival and
growth, being always higher on genuine gypsum
soils (Figure 2). This suggest that at least for this
gypsophyte, there is an intermediate strategy: it
135
primary refuges on this edaphic islands because its
capability to surpass gypsum crusts at the emer-
gence stage but also has evolved adaptive strate-
gies to perform better on such soils, as it occurs in
the case of calcicole plants. Note that under dou-
ble irrigation, recruitment does not appear to be
improved. Even more, seedlings do present sig-
nificantly lower survival and growth in calcareous
soils under double irrigation (see the significant
water*soil substratum interaction).
In conclusion our results confirm our hypothesis
that gypsum edaphic specialists base their behav-
iour in a great extent on the ability to surpass
extremely hard gypsum surface crusts (see Meyer
1986), although this seems a marginal trait as
shown by the capability to grow on a complete
array of soils and the negative effect of the crust
along the earlier development life stages of the two
studied gypsophytes.
Acknowledgements
We thank Luis G. Benavides and Dr. Albert
(‘Pizca’)for collaboration during fieldwork and
Francisco Moreno for providing valuable data.
Dr. Albert kindly read a draft version of the paper.
Financial support was provided by a MCYT
research project to A.E (REN2000-0254-P4-03).
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