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Biodiversity and Conservation 13: 1797–1808, 2004.

# 2004 Kluwer Academic Publishers. Printed in the Netherlands.

Gypsicolous flora, conservation and restoration of

quarries in the southeast of the Iberian Peninsula


and M.E. MERLO
Dpto. Biologı́a Vegetal y Ecologı́a, Universidad de Almerı́a, Ctra. Sacramento s/n, La Cañada de San
Urbano, E-04120 Almerı́a, Spain; *Author for correspondence (e-mail: jmota@ual.es; fax: +34-950-

Received 7 November 2002; accepted in revised form 30 June 2003

Key words: Ecological restoration, Endangered species, Endemic species, Gypsophytes, Gypsum

Abstract. The southeast of the Iberian Peninsula, more precisely, the province of Almerı́a, has the most
important gypsum quarries in Spain. Despite the severe environmental impact of these quarries, so far
little attention has been paid to either the effect on the flora, especially on the more restricted endemic
taxa, or to the restoration of the habitats after the end of exploitation. It is our view that restoration of
gypsum quarries should not only aim at correcting the visual impact of mining on the landscape, but
should also attend to the biological impact of the quarrying. Although the ‘cicatrising’ potential of
gypsophytes is already well known, not all the species are able to recolonise worked-out quarries. Using
species abundances, all analyses clearly show that artificial rehabilitation results in very altered com-
munities. The preservation of the gypsum flora merits prioritary protective strategies which, at least to
some extent, would make gypsum mining and nature conservation compatible.


The close relationship between plants and a particular geological substrate has long
been recognised (Rivas Goday 1969; Kruckeberg 1984). In recent times, a number of
botanists and plant ecologists have dealt extensively with this topic (e.g., Kruckeberg
1991; Roberts and Proctor 1992; Mota et al. 1993). Gypsum substrates, for example,
when occurring in dry areas, have significant geobotanical effects (Johnston 1941;
Parsons 1976; Turner and Powell 1979; Meyer 1986). Although among gypsophytes,
that is, plants inhabiting only gypsum soils, the adaptive features are not remarkable,
many of them show foliage succulence, salt excretion (Duvigneaud and Denaeyer de
Smet 1966, 1968), or deeply penetrating roots (Guerrero 1998).
Plant gypsophily has been explained in two ways. While some researchers
postulate physical constraints to account for gypsophily, others stress chemical or
nutritional requirements as causative factors (Parsons 1976; Merlo et al. 1998). On
the one hand, crusts either of biogeneous (mostly lichens) or other origin (rocky
edaphic horizons) and the hydric performance of the superficial soil layer (Meyer
and Garcı́a Moya 1989; Meyer et al. 1992) have been quoted as physical factors
inducing gypsophily. On the other hand, some of the chemical hypotheses aiming at
explaining gypsophily are deficiency of macronutrients (N, K and P), excess of
others (Ca, Mg and S), ionic antagonisms (Ca/Mg) and toxicity caused by micro-

nutrients. These restrictive factors can affect plant development in different stages
of their life cycle, for example, during germination (Merlo et al. 1997) or during
establishment (Escudero et al. 1999, 2000).
The Iberian flora has about 30 endemic gypsophytes (Merlo et al. 1998). This
enormous amount of endemic taxa is mostly due to the impressive gypsum deposits in
Spain, with over 30,000 km2 some of the largest in Europe (Riba and Macau 1962;
Ayala et al. 1986) and an antiquity which ranges from the Triassic to the Quaternary
period. Many of the vascular plants inhabiting these deposits are protected either by
international, national or regional laws (Pintos and Rodrı́guez-Hiraldo 1994). Gómez-
Campo’s catalogue (1987) lists up to 12 gypsophytes which are also mentioned in
Appendix II of the EU Habitats Directive (Anonymous 1992). Although some of these
species are also protected by the Bern Convention (Anonymous 1979), their extent of
occurrence and the area of occupancy have diminished. As a matter of fact, Blanca et al.
(1999) report a reduction of almost 15%, which continues and will presumably do so
until the expiration of the exploitation licences. This is probably the main reason why
the endemic taxa of Almerı́a, Teucrium turredanum and Helianthemum alypoides, are
also mentioned in the recently published ‘Lista Roja Española’ (VV.AA. 2000) to-
gether with Teucrium balthazaris Sennen, another endemic gypsophyte of the south-
east of the Iberian Peninsula.
The richness and rarity of the vascular flora colonising these gypsum deposits
strongly justify the official conservation of these habitats (indeed, the EU Habitats
Directive gives priority to their conservation). The future Nature 2000 Network
provides a good opportunity to promote the preservation of these gypsum outcrops
which, due to their geographically punctuated distribution, present such an extra-
ordinarily varied flora. The gypsum deposits are also the habitat of other plant
communities of great interest, such as the therophyte grasslands (e.g., Izco 1976)
and the crusts of bryolichens (Casares-Porcel and Gutiérrez 1995).
The rich variety of vascular plants and cryptogams is most remarkable in the
south of the Iberian Peninsula, particularly in the province of Almerı́a (Guerra et al.
1995; Mota et al. 1998). This province also includes one of the most valuable
karstic systems on gypseous soil in the Mediterranean area (Calaforra and Pulido-
Bosch 1997). Since these areas concentrate 25% of the gypsum extraction in Spain
(Junta de Andalucı́a 1999), the conflict between conservation and exploitation is
inevitable. The industrial use of gypsum, either as a raw material for building or as
an ingredient of stucco and cement, seriously endangers the conservation of these
plant communities and ignores their biological and evolutive value (Parsons 1976;
Meyer 1986). Gypsum quarries do indeed provide a vital and multirole material in
any industrialised society, but at the same time the environment is seriously affected
(Davis 1982). Gypsum quarrying has a tremendous visual impact on karstic land-
scapes, altering them as dramatically as limestone quarrying has done in other
places (e.g., Gagen et al. 1993; Gunn 1993; Gunn and Bailey 1993). Modern
exploitation techniques, with sophisticated machinery and more powerful
explosives, produce long-lasting alterations in the landscape. In addition, in gypsum
quarrying these effects are not only visual (pollution occurs only to a minor extent),
but they also affect the karstic system and the flora too.

Nowadays, there is broad scientific consensus on the subject of the dramatic

decline of biodiversity (e.g. WCMC 1992; Heywood 1995). This concern makes it
essential to accurately evaluate the effects of any eventual extractive activity on the
valuable flora which colonises these gypsum deposits. If it is proved that quarrying
seriously endangers plant biodiversity, then two possible environmental policies can
be implemented: in sites still unaltered by mining, native vegetation should be
preserved from any devastating effect caused by mineral exploitation; in already
altered sites, it would be advisable to try to restore the native communities, that is,
the ecological integrity previous to extraction, once mining comes to an end. As
Higgs (1997) states, ecological fidelity implies structural/compositional replication,
functional success and durability.
Spanish laws envisage the restoration of any natural landscape affected by open-
air mining (Real Decreto 2994/1982). At present, mining is not allowed if there is
no official plan to prevent or at least diminish the deleterious effects of the in-
dustrial activities. However, Spanish environmental policy does not promote the
ecological restoration of the original plant communities on gypsum deposits once
extraction is over. In fact, to our knowledge, the infrequent restoration programmes
already carried out overlook the peculiarities of gypsum outcrops and their flora.
Once more, due to the convergence of important conservationist interests and in-
tensive mining, the Iberian southeast becomes a Mediterranean hotspot where an
outstanding richness of species, many of them endemic, coexists with highly me-
nacing agents (cf. Mota et al. 1996), a situation particularly useful for observing
and assessing the conflict between development and biodiversity preservation.
In this study our aim is twofold: first, to assess the impact of mining on gypsi-
colous endemic flora and to measure the autoregenerative potential; second, to
evaluate the effects of the current artificial restoration of habitats.

Material and methods

Area under study

This work deals with the gypsum outcrops of the province of Almerı́a (Iberian SE)
distributed in three large geographical units: the depression of Rı́o de Aguas, the
corridor of Tabernas and the basin of Rı́o Almanzora. Figure 1 shows the UTM quadrats
in which both the gypsum outcrops colonised by gypsicolous flora and the location of
the quarries under study are shown. Table 1 presents some basic data of the spots.

Data analysis

Impact on flora
There is no doubt about the serious impact of mining on gypsicolous flora, parti-
cularly on strict gypsophytes of a comparatively restricted geographical distribution
(narrow endemism). However, some studies have revealed that Gypsophila

Figure 1. Location of the areas under study. The map shows the UTM 10 km  10 km grid encom-
passing the 1 km  1 km grid mentioned in the text.

Table 1. Basic data of the spots: UTM quadrats, abbreviations of the quarries sampled in each area and
gypsophytes of the non-disturbed scrubland.
Study area UTM Abbreviation Gypsophytes of the nearest
non-disturbed scrubland

Almanzora-Topares 30SWG8929 CAQI T. balthazaris (Teu_bal); Jurinea pinnata

30SWG9135 CAQI (Jur_pin); Ononis tridentata (Ono_tri);
30SWG9934 CAQII Santolina viscosa (San_vis); Helianthemum
30SWG8440 CAQII squamatum (Hel_squ); Launaea fragilis
30SWG8432 CAQII (Lau_fra); G. struthium (Gyp_str)
Sorbas-Rı́o de Aguas 30SWG8004 SOQI T. turredanum (Teu_tur); H. alypoides
30SWG8498 SOQI (Hel_aly); O. tridentata (Ono_tri); Coris
30SWG8310 SOQII hispanica (Cor_his); Chaenorrhinum
30SWG8409 SOQII grandiflorum subsp. grandiflorum
30SWG8309 SOQII (Cha_gra); S. viscosa (San_vis);
30SWF8498 SOQII H. squamatum (Hel_squ);
L. fragilis (Lau_fra); G. struthium (Gyp_str)
Tabernas-Venta 30SWG6204 VYQI Frankenia thymifolia (Fra_thy); Lepidium
de los Yesos 30SWG6204 VYQI subulatum (Lep_sub); Rosmarinus
30SWG6203 VYQII eryocalix (Ros_eri); C. hispanica
30SWG6304 VYQII (Cor_his); C. grandiflorum subsp.
30SWG6203 VYQII grandiflorum (Cha_gra); S. viscosa
30SWG6304 VYQII (San_vis); H. squamatum (Hel_squ);
30SWG6204 VYQII L. fragilis (Lau_fra); G. struthium (Gyp_str)

struthium L. cannot only recolonise worked-out quarries, but also prosper and form
populations larger than those of unaltered areas (Merlo et al. 1997, 1998). In order
to assess the impact of mining on the whole gypsicolous flora, we recorded the

abundance of species on gypsum deposits and compared the data obtained in

quarries with those of unaltered gypsum outcrops. Our work treats independently
each of the quarries located in each of the three gypsum areas of the province of
Almerı́a as defined by Mota et al. (1997). Thus, we studied six quarries in Sorbas-
Rı́o Aguas, eight in Venta de los Yesos-Tabernas and five in the Almanzora valley.
Each quarry was ascribed a sample of the neighbouring gypsicolous scrub occur-
ring in the unaltered area of the outcrop. To estimate the abundance of the species
we randomly sampled five 5  5 m2 squares. In all the squares, the cover area of the
scrubland species was recorded. Quadrat size was determined following Greig-
Smith (1983), who advises against the use of square sizes encompassing many
species with frequency rates near 100%. The time since the end of exploitation
varied from 10 to over 40 years, which led us to classify them into two categories or
phases, namely: recently abandoned quarries (20 years or less since the end of
exploitation) and quarries abandoned long ago (i.e., over 25 years). From now on,
the former will be referred to as initial phase quarries (QI), whereas the latter will
be referred to as advanced phase quarries (QII), labels which aim at depicting the
theoretical state of the autogeneous primary succession. Although it was already
known that some species, such as G. struthium, only need 10 years or even less to
profusely recolonise quarries (Merlo et al. 1998), we tried to determine how general
a pattern this was among gypsophytes. All the field sampling took place in 2001.
Some of the figures show a threefold classification of species: (i) strict gypsophytes,
that is, characteristic of the Gypsophiletalia order, or of some of its suballiances, as
suggested by Loidi and Costa (1997); (ii) ruderal species; and (iii) non-gypsophytes.
Dittrichia viscosa (L.) Greuter, Piptatherum miliaceum (L.) Coss., Artemisia bar-
relieri Besser, Helichrysum italicum (Roth) G. Don subsp. serotinum (Boiss.) P.
Fourn., Andryala ragusina L., Launaea arborescens (Batt.) Murb. and Phagnalon
saxatile (L.) Cass. were classified as ‘ruderal’ plants. These species characterise the
phytosociological units peculiar to the first-colonizing nitrophilous vegetation of
the Iberian southeast (Alcaraz et al. 1991; Peinado et al. 1992).
Since the species cover rates were not parametric, not even after their conversion
in order to assess whether or not differences were significant, we implemented a
Mann–Whitney test (Sokal and Rohlf 1995; Zar 1996). This approach made it
possible to compare the abundance rates of each of the gypsophytes between the
four habitats (M-QI, M-QII, QI-QII and, only in Sorbas, M-Re, QI-Re and QII-Re).

Effects of man-induced restoration

Despite Spanish legislation, which makes the restoration of quarries after the end of
exploitation obligatory, restored gypsum quarries (Re) are hard to find. This is prob-
ably due to the fact that some of these bills were passed before the end of exploi-
tation of many small mines and also to the absence so far of appropriate restoration
programmes for the enormous exploitations still working. As far as Rı́o de Aguas is
concerned, 10 years ago there was an attempt to restore five plots of variable area in
a quarry belonging to the Compañı́a Minera Fuente del Peral. The restoration was
limited to the covering of the quarry square with top soil. Following the above
mentioned procedure, five samples of each of the five artificially restored plots were

taken in order to compare the abundance of the gypsophytes with the corresponding
records of both the autogeneous primary succession and of the bushes of the
neighbouring, unaltered outcrop.

Results and discussion

Impact on flora
Most gypsophytes show significant differences in their cover rates, both inside and
outside the quarry (Table 2). The abundance rates of the main species can be seen in
the histograms of Figures 2 and 3, and the result of the comparative analysis of
medians for each two pairs of phases in Table 2.
G. struthium is the only gypsophyte whose cover rate in quarries is always higher
than in unaltered areas. This pattern can be seen in all the quarries and is even more
noticeable in the initial phases of the colonising process (Figure 2). There is
probably no easy explanation for this. Presumably, biological features both in
connection with the colonising potential and the endurance of G. struthium on
crystalline gypsum substrates are in some way accountable (Mota et al. 2003). The
quarry squares are extremely adverse environments and are almost nothing but
crystalline and very hard gypsum rock, that is, solid rock.
Leaving aside G. struthium, the rest of the gypsophytes show a completely
different pattern: many of them hardly occur in recently abandoned quarries or
are entirely absent. In the quarries of Rı́o de Aguas, almost all species are absent
in the first stage of succession (Figure 3a and Table 2); this also goes for the
local endemic species T. turredanum and H. alypoides. Only the pattern of L.
subulatum in the earlier stages of succession is similar to that of G. struthium
(Figure 3b).
Although in the quarries of Cuevas del Almanzora, during the earlier stages of
succession, the pattern is very similar to the one observed in Rı́o de Aguas and
Venta de los Yesos, there are some noteworthy differences, particularly if we take
into account the abundance of G. struthium (Figure 2). Firstly, the local endemic
species T. balthazaris is relatively abundant from the very beginning (Figure 3c).
Secondly, S. viscosa, an endemic plant of the Iberian southeast occurring in all
the areas under study, thrives more rapidly in the quarries of the river Almanzora
than elsewhere (Figure 3b and c). Why should this be? The physical profile of
gypsum, mostly hard and macrocrystalline in the outcrops of Rı́o de Aguas and
Venta de los Yesos, whereas it is soft and saccharoidal in the Almanzora river
valley (Calaforra and Pulido-Bosch 1999), has probably much to do with this
performance. These differences are so remarkable that gypsum karstic landscapes
with singular surface formations, such as tumuli and dolines or underground
caves, can be found only in the first gypsum type (Calaforra and Pulido Bosch
As far as the late stages of succession are concerned, most of the gypsophytes
tend to increase their coverage, although the records are well below the
values measured in the scrub (Figure 3a–c). Nevertheless, when the records of

Table 2. Results of a Mann–Whitney test to compare the cover medians of each species in the four
environments studied (M, QII, QI and Re). The sign ** indicates different medians with p < 0.01, and *
with p < 0.05. A sign indicates that the species is absent in either of the two environments being

Gyp_str n.s. ** ** ** ** **
Hel_aly – – ** – n.s. **
Teu_tur – – ** – n.s. **
San_vis – – * – ** **
Cor_his n.s. ** ** n.s. n.s. **
Hel_squ n.s. ** ** – – –
Lau_fra – – ** – n.s. **
Gyp_str ** ** n.s.
San_vis ** ** **
Cor_his n.s. ** **
Lep_sub ** n.s. **
Fra_thy – – **
Hel_squ ** ** **
Lau_fra ** ** **
Gyp_str * n.s. n.s.
Teu_bal * ** *
San_vis n.s. n.s. *
Cor_his – – –
Hel_squ * ** **
Lau_fra – – n.s.

Figure 2. Cover (%) of G. struthium in the abandoned quarries and in the non-altered scrubland in each
studied area. The cover in the restoration made in Sorbas is also shown.

Figure 3. Cover (%) of the main gypsophytes in each successional stage: (a) Sorbas; (b) Venta de los
Yesos-Tabernas; and (c) Almanzora valley.

recently abandoned quarries (QI) are compared with those of quarries abandoned
long ago (QII), all the species, even those with low colonising potential, such as
F. thymifolia, H. squamatum or C. hispanica, clearly tend to recuperate as the

Figure 4. Average cover (of the studied areas) of all gypsophytes (GIPSO-T), G. struthium (GS), all
gypsophytes except G. struthium (GIPSO-P), the non-gypsophytes (NGIPSO), and the ruderal species.
The cover in the restoration exclusively accounts for Sorbas.

autogenic primary succession proceeds (Glenn-Lewin et al. 1992). This per-

formance is in line with the general pattern already observed whereby, in the
autogeneous succession process, new vegetation tends to eventually resemble the
neighbouring vegetation (Prach et al. 2001). This pattern could be even more
intense in arid zones and may be the reason why some researchers have considered
it as a case of autosuccession (Shreve 1942).

Effects of man-induced restoration

The abundance values of gypsophytes in restored environments are considerably
lower than the records obtained in unexploited areas. Quite significantly, this also
goes for G. struthium, in contrast with the pattern observed in the primary suc-
cession (Figure 4). However, we must interpret this fact with caution, since re-
storation is relatively recent (about 10 years). Nevertheless, the meagre role played
by G. struthium can be interpreted as a result of the reduced invading potential of
gypsophytes in this type of habitat. On the other hand, the high invading potential
of ruderal anemochorous species, especially D. viscosa and A. barrelieri, in man-
induced restorations is very impressive. This pattern strongly contrasts with the
absence of these species in quarry squares where no man-induced restoration has
ever taken place (Figure 4). Bearing in mind the nature of the soils employed in
artificial restorations, this performance would support the suggestion already made
by Johnston (1941) that even a fine clay layer may impede the growth of gypso-
phytes, however gypseous the soil underneath may be. This fact not only highlights
the peculiarities of gypseous soils, but also alerts us to the deleterious effects that
restorations disrupting the natural soil profile can have on the preservation of these


1. There is no doubt of the serious impact of mining on the gypsicolous flora,

especially the comparatively more restricted endemic taxa. H. alypoides and T.
turredanum are the epitome of this vulnerability. The restoration of their original
populations should be strongly promoted after the end of exploitation.
2. Keeping in mind that gypsum outcrops are considered prioritary habitats for the
EU and the extracting centres in Spain tend to be concentrated on the most
biologically valuable sites, restoration should aim particularly at the ecological
recovery of the original environments.
3. In this respect, restoration strategies should rely on the high colonising potential
of some gypsophytes, particularly of G. struthium, but at the same time not
overlook the use of other species. Restoration programmes of gypsum quarries
must be an opportunity to settle sustainable populations both of the more strictly
endemic plants and also of very rare species (e.g., R. eriocalyx).
4. As a whole, gypsophytes clearly tend to recover their former populations in an
autogeneous primary succession process, a fact which reveals the extraordinary
resilience of these habitats. The almost total absence of invading plants is one of
the most remarkable features of the succession on gypsum soils. This is pre-
sumably due to the fact that, after the end of exploitation, the quarry square is a
hard and almost barren surface of crystalline gypsum, hardly suitable for the
settlement of plants, if they are not extremely stress-resistant species.
5. The widely used technique of covering the quarry squares after the end of ex-
ploitation with top soil seems to promote vegetal communities very different from
the aboriginal gypsicolous ones. Due to their peculiar ecological profile, gypseous
soils should be restored with species and procedures especially tailored not to
‘bury’ the gypsum under layers of soil of a very different edaphic composition.

To sum up, the rehabilitation of gypsum quarries should not only aim at correcting
the visual impact of mining on the landscape, but should also concern itself with the
biological impact of the quarrying. The ever-growing quarrying area seriously
reduces the sites suitable for the endemic flora which colonises them. As gypso-
phytes inhabit edaphic archipelagos, we face the risks and eventually fatal con-
sequences of destroying and fragmenting natural habitats.


This research was financially supported by a grant from the DGES (PB98-1385).


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