You are on page 1of 22

Biodiversity and Conservation 12: 18611882, 2003.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

Rarity and threat relationships in the conservation


planning of Iberian flora

FELIPE DOMINGUEZ
LOZANO 1, *, JUAN CARLOS MORENO SAIZ 2 and
HELIOS SAINZ OLLERO 2
1

Department of Environmental Science and Policy, University of California, One Shields Avenue, Davis,
2
( Botanica

), Facultad de Ciencias, Universidad Autonoma


CA 95616, USA; Departamento de Biologa
de Madrid, E-28049 Madrid, Spain; * Author for correspondence (e-mail: fdominguez@ ucdavis.edu;
fax: 11 -530 -752 -3350)
Received 14 March 2002; accepted in revised form 27 September 2002

Key words: Iberian peninsula, Management activities, Non-parametric principal component analysis,
Plant conservation, Rarity, Threat
Abstract. We analysed the threatened flora of Iberia (including the Balearic Islands) in order to define and
explain factors related to levels of rarity and threat. Conservation measures were derived from the
relationships observed. We used a random sample of 59 narrowly distributed plant species from the pool
(588 species) of potentially endangered flora. Twelve variables were used to classify species into groups
based on a multivariate technique: non-parametric principal component analysis. Our results do not
indicate a single management model driven by a single mechanism of rarity. Four classes of rare plants
were produced from the statistical algorithm: agamospermic species, plants associated with water,
endemics, and range-margin (geographical-limit) plants. Some specific strategies for each of these groups
are proposed, allowing further discussion and assessment. The overall pattern in conservation practice of
threatened Iberian plants seems to be defined by three of the variables in use: ecological specificity,
geographical rarity and rate of threat. None of the biological variables in the sample show particularly
strong trends in the data.

Introduction
There are several approaches to the conservation of threatened plants. Typical
conservation management actions or plans are developed as emergency responses to
crises and are developed in isolation and from idiosyncratic points of view
(depending on whether managers or scientists are involved). Despite the obvious
necessity of specific species approaches, there is also a need for a broad focus in
plant conservation.
Synthetic perspectives for conservation planning have been proposed, but with a
variety of aims and objectives (e.g. Toledo 1993; Schemske et al. 1994; Morse 1996;

Palmer 1996; and for Spain: Banares


Baudet and de Benito 1990; Gomez
Campo
Zarzoso 1994). Most of these works rely on the quality and origin of
1990; Simon
information used and often face an irremediable bias from the selection of species
under study. One of the most active fields is the development of a species selection
process using non-subjective criteria (Carroll et al. 1996; Palmer et al. 1997; de
Lange and Norton 1998; Keith 1998). Verifying the threat (i.e., a quality that

1862
decreases the evolutionary potential of the plant) is not easy for a number of reasons.
First, time causes uncertainty; there are lagged responses of species to perturbations.
Second, the unit that is threatened might not be clear; the individual, the population
and the taxonomic species are all possible objects of threats. Third, and perhaps
most important of all, most cases lack objective data documenting habitat or
population reductions, genetic constraints, or competitive viability. The assessment
of threat easily becomes a matter of judgement and therefore subjective. As a first
step to avoid potential misinterpretation, stenocory (narrow range) has been widely
employed in studies dealing with the process of selection of threatened flora at
different geographical scales (e.g. Ayensu and DeFilipps 1981; Barreno et al. 1984;
Dixon 1994; Carreras et al. 1996; Lewis 1998; Valencia et al. 2000). Following this
approach, we use data on range size, which can be relatively affordable to collect, as
indicator of putative endangerment.
Assuming rarity is a valid predictor of conservation priority, the objectives of this
study were fourfold. The first goal was the characterisation of shared properties and
differences of the rare plants of Iberia. Second, we analysed the importance of
environmental, life history, anthropogenic, and biogeographical factors related to
perceived endangerment and conservation priority status. Third, patterns among rare
taxa that emerge from this analysis are used to offer management recommendations
to improve conservation planning in the Iberian Peninsula. Finally, we try to identify
key attributes and processes that we predict will likely be related to conservation
prioritisation.
Some of the novelty of this study lies in the application of a statistical analysis to
conservation prioritisation. Our methods are inspired by quantitative biogeography,
where multivariate techniques have a long tradition (see Birks 1987). Although the
use of ordination and cluster analysis in conservation is not new (e.g., Given and
Norton 1993; Hall 1993), our study uses a non-parametric, multivariate, qualitative
ordination (PRINCALS; Gifi 1991) that has been little used in ecology.

Materials and methods


A broad definition of the Iberian Peninsula is used for the present work, including
Berlengas...), the
the main continental part, the closer islets and rocks (Alboran,
entire Pyrenees (considering the French part) and the Balearic archipelago. Over this
territory, the conservation status of the vascular plants present in the area naturally
or putatively naturalised has been revised.
As a first step, we reviewed the Iberian floristic literature up through 1997,
gathering information from 375 references. The purpose was to produce a preliminary list of rare taxa extracted from the whole flora. Species were chosen exclusively
on geographical rarity criteria. Unfortunately, there are very few data to assess other
types of rarity (abundance, population density or number of populations) in the
Iberian literature. The criteria we used to make the selection of rare species are in
accordance with two of the types of rarity sensu Rabinowitz (1981). These are:
1. Species with narrow distribution: 10 or fewer UTM-grid squares of 10 3 10 km.

1863
Although the adoption of a precise limit to quantify rarity in chorology is always
arbitrary and area dependent, similar figures have been set up in other classifications of threatened plants (Perring and Farrell 1983; Briggs and Leigh 1996;
Palmer 1996).
2. Species with wider distributions but specific habitat requirements. As above, a
convention limit has been chosen: species reported in fewer than 20 UTM-grid
squares. In this way, plants from steppe habitats, extrazonal environments (e.g.,
bogs, lagoons and saline habitats, vernal pools, sand dunes), species associated
with scattered and infrequent substrates in Iberia (gypsum, dolomite), or from
isolated disturbed places could be included.
Applying these criteria, 588 taxa were identified (a complete list appears in

Domnguez
Lozano (2000), p. 331). These plants represent a well-known and
homogeneous group with precise distribution and taxonomic status and are validated
by a majority of the botanic references consulted.
To complete this analysis, the list was sorted alphabetically and 1 of every 10 taxa
was selected. The resulting sample of 59 plants (Appendix 1) is the object of the
statistical analysis presented in this paper.
Properties of the descriptors and statistical treatment
Available information about different species attributes was gathered for the
selected taxa. Each species was scored using 12 variables or descriptors (see Table

1, and for a more detailed description: Domnguez


Lozano (2000) op. cit., p. 267).
Principal component analysis by means of alternating least squares (PRINCALS)
was applied to this data set. The main difference with conventional principal
component analysis consists in the variable transformation utility that allows the use
of not only numeric but also qualitative data (Gifi (1991), p. 177). Species attributes
are scored in categories which can be ordinal or not, acquiring different measure
levels (ordinal, binomial, single nominal and multiple nominal), each of them with a
specific statistical treatment given in the test. This option allows a more exploratory
and flexible approach in the treatment of heterogeneous data coming from different
sources. It is one of the main advantages of the technique for the data set used in this
work, where the nature of the categories of the 12 variables was diverse.
One concern for this method is that too much significance may be given to
categories with very low absolute frequencies. This problem is particularly likely
among multiple nominal variables (Kroonenberg et al. 1997). In order to avoid this,
we recodified some of the initial variables, grouping poorly represented categories
(see Appendix 2). The PRINCALS option was run using the SPSS statistical
software package (SPSS Inc. (1994), p. 49).

Results
The PRINCALS algorithm calculates parameters in a similar manner to other

1864
Table 1. Variables and categories (identification labels used in later statistics appear in brackets).
Variable

Categories

EFFECT OF ANTHROPOGENIC INTERACTION

1. Positive (pos)
2. Not detected (ndet)
3. Habitat fragmentation, direct population decrease or genetic
alteration (1effe)
4. Whichever combination of two of the processes mentioned
above (2effe)
5. Sum of effects of the three processes (3effe)
1. Without conservation interest (nint)
2. Considered interesting by academia or legally protected or, at
least in part, included in a protected area (prot)
3. Legally protected and also, at least in part, included in a
protected area (maxpro)
1. Classical: until the end of the XIXth century (clas)
2. Modern: from 1900 to 1980 (mod)
3. Actual: since 1980 (act)
1. Null: without registered synonymy (nul)
2. Low: one or two synonymous (l)
3. Medium: three (m)
4. High: four or more (h)
1. Annual (ann)
2. Biennial (bien)
3. Until 5 years (,5)
4. More than 5 years (.5)
1. Winded anemochorous (wind)
2. Villous anemochorous (villo)
3. Semachorous (sema)
4. Autochorous (auto)
5. Endozoochorous (endo)
6. Exozoochorous (exo)
7. Myrmecochorous (myrme)
8. Unknown (unkn)
1. Entomophylous (ento)
2. Anemophylous (ane)
3. Absence (abse)
1. Yes (y)
2. No (n)
3. No data (ndat)
1. Facultative allogamous (facalo)
2. Strict allogamous (stralo)
3. Facultative autogamous (facauto)
4. Agamospermous (aga)
5. No information (ninfo)
1. Calcarious (calc)
2. Acid (acid)
3. Gypsum (gyps)
4. Serpentine (serp)
5. Indifferent (indi)
6. Saline (sal)
7. Imprecise (impr)

LEVEL OF PROTECTION

TIME OF DESCRIPTION

RATE OF TAXONOMIC
SYNONYMY

LIFE CYCLE

TYPE OF DISPERSION UNIT

TYPE OF POLLINATION

CLONAL GROWTH

TYPE OF BREEDING SYSTEM

SUBSTRATE

1865
Table 1. (continued)
Variable

Categories

HABITAT

1. Woodland communities (wood)


2. Sandy and stony poor soils, cliff platforms (sand)
3. Weedy associations and high perturbance places (weed)
4. Rock walls and crevices (rock)
5. Shrublands (shru)
6. Halophytic lagoon shores (lago)
7. Clears and woodlands fringes (clea)
8. Bogs and springs (bog)
9. Mobile screes (scree)
10. Unidentified (unind)
0. Distribution limit (limi)
1. Endemic (end)

BIOGEOGRAPHY

Table 2. Analysis values for the sample of 59 taxa (names of recodified variables appear in lower case
letters).
Variable

TYPE OF POLLINATION
Breeding system
Substrate
TIME OF DESCRIPTION
Dispersion units
Habitat2 a
BIOGEOGRAPHY
LEVEL OF PROTECTION
CLONAL GROWTH
Anthropogenic interaction
Life cycle
RATE OF TAXONOMIC SYNONYMY
Habitat1 a
Eigenvalue

Component loadings
Dimension 1

Dimension 2

20.437
20.157
20.768
20.701
0.730

0.603
20.356
0.513
20.412
20.286
0.416
0.683
0.2895

20.722
20.789
20.095
0.282
20.059
0.672
20.262
0.503
20.010
20.270
0.360
20.016

0.1844

Total variance accounted for

0.713
0.647
0.598
0.570
0.537
0.459
0.433
0.380
0.263
0.242
0.211
0.174

Absolute values over 0.6 for loadings appear in boldface. a Multiple nominal variables have independent
loadings on each axis.

multivariate analyses. Table 2 shows the factor loadings for two dimensions with 12
variables over 59 species. The eigenvalues, which reflect the reliability of the
analysis, explained 44% of the total sample variance across the first two dimensions
(PRINCALS axes). For simplicity, we restrict our attention to these two axes.
Figure 1 depicts the coordinates of the categories of each variable along the two
axes considered. Two categories emerge as the most explanatory descriptors for
dimension two (Type of pollination and Breeding system), defining an isolated
group of plants. These categories are abse (without recorded pollination) and
auto1 aga (where cases of autogamus and no data breeding systems are included).
They are accompanied by category lago (species on saline lagoon shores) of the
Habitat variable. In addition, there is another pattern of variation along the first axis,

1866
which is in agreement with the loadings values of the Substrate and Dispersion units
variables on axis I (see Table 1). Other descriptors contributing to the explanation of
the variance for this axis are Time of description, Habitat and Biogeography (with
20.701, 0.683, and 0.603 loadings values, respectively). Nevertheless, the distribution of axis scores is influenced by high quantifications on the second axis.
The graphic on the taxa scores (Figure 2) follows the same model, showing a
segregation of one group of species from the rest. Three species [Limonium
migjornense (Limmig), Hieracium vegaradanum (Hieveg), and H. glaucophylum
(Hiegla)] appear in the corner of the lower left quadrant. Each of these species is
characterised by the categories abse and auto1 aga, and they constitute the first
well-defined species group of the analysis.
Although some intermediate species lay in between, Avena eryantha because of
its auto1 aga category, and Limonium stenophyllum and Puccinelia fasciculate spp.
pungens due to their lago category, the rest of the species are better clustered by the
correlation with axis I.
This analysis is dominated by the high scores of the three-species assemblage in
the left bottom corner, and the prominent differences from the rest of the species
obscure the distribution of the main body of species on axis I, where not very much
spread of the data occurs.
For this reason, once the model showed the peculiarity of this first group
(Limonium migjornense, Hieracium vegaradanum, and H. glaucophylum), they
were removed in a second run of the multivariate correspondence analysis. The aim
was to improve the interpretation of the structure of the data of the remaining
species and variables without the presence of outlier scores.
The elimination of these species required a new recodification of the Habitat

Figure 1. Category coordinates from the variables. Not all labels have been plotted.

1867

Figure 2. Scores for the 59 taxa of the sample (not all labelled). Each species is labelled with a six- or
nine-letter abbreviation for the first three letters of the genus, species and subspecies. The dashed line
emblazes the first group of species referred to in the text.

variable and Type of pollination was transformed into binomial due to the loss of
one of the categories (Appendix 3); the rest of the variables remained the same.
The results of this new test are summarised in Table 3; the new calculated
eigenvalues point out that no significance was lost. Nevertheless, a change in the
power of discrimination of some descriptors has occurred. Type of pollination and
Breeding systems now explain little of the variance, stressing even more their
relationships with the species eliminated from this second analysis. The loss of the
abse category (pollination absence) in the Type of pollination variable due to
recodification of its very low frequency contributes to the statistical change. Habitat
retains its explanatory power; again especially on axis II. In addition, Level of
protection and Time of description become important (0.638 and 0.618 for the total
variance values). Modifications of values for the less useful variables (Rate of
synonymy, Life cycle, and Clonal growth) are small.
The second dimension reflects these changes in the values of the variables.
Habitat is the most important variable along this axis; and aqua (species from
lagoons, peat bogs and springs) possess the most distant coordinates from the origin
(see Figure 3). This tendency is reinforced by three other classes: maxprot (maximum protection), ane (anemophylous sensu lato) and 12 effe (at least two effects of
the three considered, habitat fragmentation, population decrease or genetic altera-

1868
Table 3. Component loadings, total variance accounted for, and eigenvalues for the descriptors of the
second assessment of the analysis.
Variable

LEVEL OF PROTECTION
TIME OF DESCRIPTION
a
Habitat2
BIOGEOGRAPHY
Dispersion units
Anthropogenic interaction
TYPE OF POLLINATION
Substrate
CLONAL GROWTH
Life cycle
Breeding system
RATE OF TAXONOMIC SYNONYMY
Habitat1 a
Eigenvalue

Component loadings
Dimension 1

Dimension 2

20.304
20.766

0.648
0.600
20.032
0.027
0.620
0.522
20.492
0.447
0.412
0.592
0.2567

20.738
0.204
20.824
0.147
20.283
20.639
20.631
20.003
0.038
0.024
0.079
20.125

0.1832

Total variance accounted for

0.638
0.628
0.515
0.442
0.440
0.410
0.399
0.384
0.274
0.242
0.206
0.186

Absolute loadings values over 0.6 appear in boldface. a Multiple nominal variables have independent
loadings on each axis.

tion). However, two other descriptors now contribute to the explanation of variance:
Level of protection (0.548) and Anthropogenic interaction (0.397).
Readjustment of variance also resulted in a better segregation along axis I,
particularly for the Biogeography and Time of description descriptors (now with a

Figure 3. Category scores for descriptors of the second multivariate analysis. Some labels have been
omitted for clarity.

1869
high loading of 0.648 and 20.766, respectively), but also for others already
important in the previous test like Habitat, Unit of dispersion and Substrate.
Following this model, the species score plot (Figure 4) contains a fairly scattered
group, far from the origin and through the negative values of axis II. Distinctive
species are Puccinellia fasciculata subsp. pungens (pucfaspun), Festuca frigida
(fesfri), Centaurium somedanum (censom), Delphinium montanum (delmon) and
Senecio coincyi (sencoi), all belonging to the aqua recodified category of the Habitat
variable (Figure 5a) and maxprot or prot of level of protection. All of these species,
except Festuca frigida, have been reported to respond negatively to human disturbances (12 effe and 1 effe categories). This group of species also contains two of the
three anemophylous species of the sample. This new assemblage of species forms
the second rare species group derived from the analysis characterised for its affinity
to aquatic environments.
In addition, the upper part of the plot also shows some segregation of species. The
margin distribution species (limi) and those described in classical time (clas) have
high axis I scores, outlining the third group of this analysis. The fourth and last
group, endemics, lies on the left part of axis I (category end of the Biogeography
variable, see Figure 5b). This group is also distinguished by scores for the Habitat

Figure 4. Species scores for the second PRINCALS analysis of the remaining 56 taxa (not all the scores
are labelled). The dashed line distinguishes the second group of plants from the rest.

1870

Figure 5. Species scores for the second PRINCALS analysis. In this case they have been labelled using
the categories of the most significant variables of the test to show the segregation of species according to:
(a) Habitat variable, and (b) BIOGEOGRAPHY variable.

(Figure 5a) and Substrate variables. Species associated with scree and most of the
ston and shru associates are distributed on the left side of the scatterplot and species
associated with weed, wood and some belonging to the clea categories are on the
right side, where category indi of Substrate is also found. Therefore, the variation
along the first axis of this second PRINCAL analysis was useful to differentiate two
more groups of rare plants in the sample. These groupings are driven by the
biogeography variable.

Discussion
Species assemblages
Our multivariate analysis distinguished four groups of rare species, namely agamospermic species, plants associated with water, endemics, and range-margin (geographical-limit) plants. The first group (agamospermic plants) is defined by the lack
of functional pollination and by an asexual breeding system. Species from our data
set include Limonium migjornense, Hieracium vegaradanum, and H. glaucophylum.
This group of plants is similarly characterised by habitat, fragmented and non-stable
places range from saline ponds, to open or altered spaces, and woodland clearings,
respectively. These environments seem to sustain higher proportions of agamospermic species and the processes that favour them (polyploidy) (Grant (1989), p. 336).
Fragmented and degraded habitats are often associated with stresses such as extreme
microclimatic conditions or high frequency of perturbations. Richards (1997, p.

1871
400) suggested that agamospermic species have higher fitness than their sexual
counterparts where such disturbances of physiologic equilibrium occur. In addition,
some phenomena contributing to persistence of polyploids, such as low population
numbers, are common in differentiated habitats (Thompson and Lumaret 1992). The
species identified in our analysis indeed have those properties.
Others have suggested considering agamospermic species as a group for conservation purposes (Holsinger and Gottlieb 1991; Skinner et al. 1995; Morse 1996;
Hooper 1998). Despite of this, our analysis shows no association present between
the group and level of protection and anthropogenic interaction variables.
We conclude that it may be important to consider the protection status of
agamospermic species in the Iberian peninsula. Likewise, it is useful to consider that
the vegetation landscape of the territory under study has experienced a long history
1985; Garca
Anton
et al. 2002). If
of anthropogenic impacts (e.g. Pons and Quezel
the relationship between agamospermy and unstable, disturbed habitats is consistent, then we might expect that fragmentation, either natural or man-made, may
promote the appearance of polyploid complexes and therefore agamospermy.
In the case of naturally fragmented habitats, e.g. saline ponds or rocky places,
management should take into account both sexual species and asexual complexes.
The natural interchange (immigration and emigration) between populations of the
former should be preserved, avoiding artificial alteration.
In the case of anthropogenically fragmented habitats (clearings, forestry roads,
fire lines, quarries) these disturbances may have promoted agamospermic establishment to the detriment of sexual relatives less suited to fragmentation. In this case,
identification and location of original sexual species should be a conservation
priority. When reintroducing new populations of sexual relatives of agamospermic
species, sites should be carefully chosen to assure no contact. It is also advisable to
avoid potential dispersal mechanisms that could increase possibilities of introgressions among asexual lineages.
In summary, we recommend an assessment of the natural evolutionary potential
of agamospermic species among Iberian plants for conservation purposes. This
recommendation is supported by information from other specific cases (Palacios and

Gonzalez-Candelas
1997) and review articles (Whitman et al. 1991; Ranker and
Arft 1994) that suggest the biological processes associated with agamospermy,
polyploidy and hybridisation should be part of conservation policies. Doing this
could be part of a wider approach in relation to the species concept debate in this
matter (Rojas 1992; Soltis and Gitzendanner 1999). Clarification of phylogenetic
relationships and differentiation between sexual and asexual species will help to
establish the best protection methods. The frequency of agamospermy is considerable, but its exact proportion has been difficult to estimate (Keith (1998) op. cit., p.
425). In our sample, asexual species represent 5% of the total. If the whole (not only
threatened) Iberian flora is considered, the figure will probably be larger and likely
to increase as knowledge of systematics, breeding systems and polyploidy levels
improves.
The second species group identified in our analysis, plants associated with water,
is grouped by ecological properties (habitat), conservation coverage (type of

1872
Table 4. Protection measures for the second group of taxa.
Taxa

Protected area

Centaurium somedanum Somiedo Natural Park


Delphinium montanum
Festuca frigida
Senecio coincyi
Puccinellia fasciculata
subsp. pungens

Sierra Nevada National Park


Natural Park Lago de Sanabria
Laguna de Gallocanta Refuge

Legal cover
Habitat Directive. Asturian Catalogue of
Threatened Species
Catalonian plan of natural interest sites
Andalucian Catalogue of protected species

Habitat Directive. Aragonian Catalogue of


Threatened Species

protection) and to a lesser extent by their rate of alteration (anthropogenic interaction). These are specialised plants living in montane peat fens and bogs (mostly
associated with springs and streams) or in saline lagoons. Both of these habitats are
extrazonal, scattered and rare in the Peninsula and contain water most of the year.
These habitats are particularly sensitive to natural fluctuations in rainfall associated
with the Mediterranean climatic region. In addition, human alterations such as the
use of springs and bogs for drinking water and agriculture development of endoreic
lagoons, are frequent added sources of perturbation (Aldasoro et al. 1996; Pedrocchi
1998, p. 80).
The second run of PRINCAL on the Iberian data shows a clear relationship
between plants associated with water and the interest-of-protection variables (see
Table 4). Most of these plants retain some populations within protected areas, and
this status secures some measure of preservation against possible direct threats. In
situ measures are complemented with legal status because the majority of these
plants are included in official catalogues of protected species.
Our results underscore the coherence of the present Spanish planning for this
assemblage, where fragile species, with strong development pressures on their
habitat, are targets for protection measures. Nevertheless, examples to the contrary
exist within the Iberian flora: Carex lainzii is an endemic of endoreic depressions
1994) that is threatened and without legal
from the northern plateau (Luceno
protection at the moment.
Finally, it is important to point out the problems associated with the biological
characteristics of aquatic plants. Although monographic chorological works exist
(Cirujano et al. 1992), the distribution of threatened aquatics is usually poorly
known, even in fairly well studied areas (Skinner et al. 1995). This lack of
information may be related to the variability and seasonality of wet Mediterranean
habitats. Sudden oscillations of population numbers are common among aquatic
plants, adding difficulties to the interpretation of demographic monitoring and to the
assessment of the level of threat in these plants.
The remainder of the species analysed in our data set are clustered in two groups
divided by endemism (Biogeography) and most of their properties are related to this
variable.
Endemic species appear to be associated with two particular environments. One is
composed of screes and rocky escarpments (walls, ledges, crevices and poor-soil
microsites) of the Iberian mountain ranges and the second is formed by eastern

1873
Mediterranean shrublands. The first relationship has already been reported for the

Iberian flora (Montserrat and Villar 1972; Gomez


Campo 1985). These habitats
could have facilitated speciation of isolated and rare taxa. In addition, these habitats
may provide an escape from competition or predation. Thus, taxa with opportunistic
strategies, such us early successional species, are most likely to succeed in these
environments, due to certain biological properties that allow them to face fragmentation and spatial randomness, and to be the object of future environmental
differentiation. From our sample, rare species restricted to isolated rocky environments may have evolved from a widely distributed sister taxa: Aquilegia pyrenaica
summit in Cazorla range) and A. vulgaris
subsp. cazorlensis (saxicolous of Cabanas
(weedy representative), Brimeura duvigneaudii and B. amethystina, Centaurea
mariana and C. ornata, Erodium paularense and Erodium glandulosum or Petrocoptis grandiflora and the genus Silene.
The second group of endemic species is associated with Mediterranean shrubland
formations (shur). These taxa belong to shrub communities from the southeastern
sector of the territory (Biscutella dufouri, Centaurea carratracensis, Helianthemum
alypoides, Teucrium subspinosum, etc.). Our identification of this group of species
supports the need for concern for this especially diverse and unique vegetation type
(Mota et al. 1996). The region represents the marginal range of some North African
shrubland communities and contains isolated habitats that are the product of a
diversity of patchily distributed substrates (Peinado et al. (1992), p. 45). It is part of
a general trend in the distribution patterns of the Iberian endemism, which increase

towards the southeast corner (Hernandez


Bermejo and Sainz Ollero 1990; Castro
Parga et al. 1996).
Examining the species protection patterns for endemics demonstrates that it has
not been overlooked by conservation action: almost the entire sample is legally
protected or included in conservation areas. Only three species do not receive any
attention: Centaurea mariana, Aristolochia bianorii, and Sideritis bubanii, but none
of these have been classified with a high category of threat in up-to-date protection
lists (AAVV 2000).
The endemics comprise the biggest assemblage of rare and protected flora (71.2%
in the sample) and without doubt are the most important conservation target.
Nonetheless, our results indicate that risks caused by human activities (Anthropogenic interaction variable) are not characteristic of this group. Of course, there are
well-known exceptions: Artemisia granatensis, Borderea chouardii, and Rosmarinus tomentosus. In the sampled list, the majority are not directly threatened.
Furthermore, cases exist where human disturbance favours new colonisation
(Genista dorycnifolia or Centaurea carratracensis). The safer conditions of some
habitats associated with high rates of endemism in Iberia are likely to explain why
these rare species tend to have little association with human disturbance. Cliffs and
walls in mountain ranges and semi-desert shrubland communities are frequently
subjected to less human interference. Thus, a non-interventionist approach would be
recommended in a general plan for conserving Iberian endemicity. The preservation
of enough habitat in each case seems to be the most successful measure. Nevertheless, because of continuous processes of environmental degradation and the lack of
suitable habitats in the territory, routine demographic monitoring and ecological

1874
surveys of the most sensitive species are recommended to assess the alteration of
natural establishment and recolonisation dynamics.
The margin-range species are the fourth and last association. This group consists
of plants with different chorological origins. Some of them have a distribution
centered in North Africa (Anthemis crysantha, Asphodelus roseus, Nepeta apullei,
etc.). Others occur in the central European mountain ranges (Cochlearia pyrenaica,
Dracocephallum austriacum, Epilobium nutans, etc.), with certain small populations in the Peninsula. The rest are Circunmediterranean (Allium commutatum,
Fumaria flabellata, Iris juncea, etc.). They grow mostly in abundant habitats:
woodlands (climax formations) and associated environments (clearings and perturbed sites). The majority are not particularly rare in the rest of their distribution
range. This could be the reason why our analysis connects these plants with the
classical period of taxonomy.
Traditionally, a lower conservation priority has been assigned to these plants, and
they are not well integrated in the protection system of Spain (nint category of
Figure 3). This is true mainly for species growing on clearings and secondarily on
degraded places: Anthemis chrysantha, Asphodelus roseus, Melilotus italica, Potentilla inclinata, and Euphorbia gaditana (although the latter is legally protected).
Few examples exist where plants representing this group have been taken into

1993). Our model


consideration in the Peninsula (Gomez
Campo and Herranz Sanz
detects a protection weakness in this case.
Descriptors order of importance
Some descriptors have been specifically useful to outline particular groups of
species in the sample, namely type of pollination and breeding system for the first
analysis and biogeography for the second. Nevertheless, habitat could be considered
one of the most important variables for the overall explanation of the data structure.
Three groups of species contain well-defined environmental preferences. It could be
said that the type of environment where a plant grows is connected not only to its
rarity but also to its threat. This proposal is not a novelty and the results are in
agreement with several works (Kruckeberg and Rabinowitz 1985; Given and Norton
1993; Fiedler 1995; de Lange and Norton 1998; Lobo et al. 2001). The Habitat
variable splits species of sensitive habitats in the Mediterranean area (lagoons,
mountain springs and peat bogs) from other plants with affinities for woodland
communities, weedy associations and clearings. Conservation concern has been
clearly identified for the first group, but not for the second. Rocky and scree habitats
and shrub communities are intermediate. These habitats are characteristic of a high
proportion of endemics and these species typically garner the top protection ranking.
The main reason to consider biological attributes in our analysis has been to test
for patterns in the endangerment of rare species. Unfortunately, the biological
descriptors we employed in this analysis do not seem to be associated with any
pattern. We conclude that their effects may be better studied with a different
approach. Differences among plants may be observable if common species were
included in the test, as some authors have suggested (Kunin and Gaston 1993; Bevill
and Louda 1999).

1875
The Pollination descriptor has discriminated agamospermous species from the
rest, since, except for wind-pollinated grasses of the sample (Avera eriantha,
Festuca frigida and Puccinellia fasciculata subsp. pungens), the others are insectpollinated. A high proportion of entomophylia in narrowly distributed plants has
previously been mentioned (Kruckeberg and Rabinowitz (1985) op. cit.). In
addition, there is a generally conspicuous representation of entomophylous pollination in Mediterranean environments (Herrera 1988). For this reason, special
consideration of plantpollinator interactions would be generally useful for Iberian
plant conservation.
On the other hand, it seems that no significant relationships between conservation
priority and reproductive specificity (Given and Norton 1993) or between rarity and
breeding system (Weller 1994) have been discovered in previous studies. We also
found no such relationship. Although in this case, a lack of definition (apart from
agamospermic) could explain this pattern.
The remaining variables (Synonymy rate, Life cycle, and Clonal growth) are not
significant. The last two, with the data available, do not define any demographic
pattern in relation to rarity and threat phenomena, but a relationship between rarity
and extended life cycles has been found in other floras (Hedge and Ellstran 1999).

Conclusions
Although the use of distributional data based on stenochory criteria is widely used to
identify conservation priorities, at present, the process of protection listing in Spain
is not free of subjective interpretation in the quality of threat. One problem in
resolving this subjectivity is that there is a lack of biological data (population trends,
colonisation abilities, fine-scale systematics) that could assist selection.
Nevertheless, building an optimum Iberian conservation strategy should take into
account a triad of attributes: environmental range (defined in this work by the
Habitat variable), level of geographic rarity (with end and fini categories of
biogeography) and rate of threat (Anthropogenic interaction and Level of protection). These descriptors summarise the concepts of specificity, originality and
fragility, needed to sustain and develop a coherent protection policy. Biological
variables, although very important for the explanation of the status of many
threatened plants, do not seem to establish any patterns among the species sampled
in this study. Thus, with these data we cannot develop generalisations about the
influence of biological variables on rarity and threat.
These results do not promote broad-scale planning. Instead, specific management
measures, in close relationship with the three factors mentioned, are likely to be
most effective. Protection of agamospermic plants (and of microspecies and neoendemism) requires a specific treatment depending on the implication of human
activities in their origin or promotion. Active measures (population management,
habitat restoration) must be applied mostly to the second group of species identified,
which are related to especially fragile and threatened habitats (streams and mountain
springs, peat bogs, endoreic lagoons, etc.). There is a less altered floristic component, the endemics, which is nonetheless very original and appreciated in

1876
conservation planning. We recommend the maintenance of biological processes
(pollination, dispersion, microenvironmental availability) related to their demography and evolutionary potential. Finally, the plants in our data set that are rare by
virtue of being at their distribution margins, although they may exhibit biological
attributes that decrease their threatened status, deserve specific protection for three
reasons. The maintenance of these range edge populations: (1) assures the representation of all the phytogeographic elements in the Iberian Peninsula, (2) maintains
within-species diversity; and (3) facilitates the potential for natural speciation.
The close relationship between habitat and species in our study highlights human
alteration of environments as one of the key elements in the threat of the Iberian rare
plants. As shown, it can influence levels of agamospermy. The anthropogenic
perturbation of particular aquatic environments also increases the vulnerability of
some of their plants, particularly those species that do not have biological attributes
that may facilitate resilience (long distant dispersion, seed dormancy, clonal
growth). Fragmentation processes could be particularly serious when rare plants
have low-density distributions or poor recolonisation and establishment (i.e. some
endemics) and their effects are difficult to appreciate in the short term. On the other
hand, human alterations can eliminate geographic barriers to reproduction among
related species, stimulating genetic interchange and degradation of pure lineages
(Colas et al. 1997). Finally, disturbance can cause rare species to become less rare in
the case of plants related to human activities.
Present conservation planning in the Iberian peninsula, not very surprisingly,
seems to be well designated in most of the cases, as shown by our multivariate
analysis. But this does not ensure complete effectiveness. The legal protection of
Iberian plants has received a strong encouragement recently, but it is still lacking a
true management infrastructure. Perhaps more time is needed before Spanish plant
conservation management moves from its current idiosyncratic and anecdotal phase.
Gathering sound knowledge about the biology of Mediterranean rare plants also
requires more time. Even with better information, it will be very complicated to
secure the long-term viability of populations and the processes that naturally
regulated species and facilitated their differentiation. Plant conservation demands a
combination of social, economic and scientific structures (Norton 1986; Grizzle
1994; Schemske et al. 1994) that at present is very difficult to maintain.

Acknowledgements
Some useful data for the species sampled in the analysis have been kindly provided

by: Maria Lurdes Serpa Carvalho (Nucleo


de Botanica,
Instituto da Conservacao da
do Ambiente e Recursos Naturais de Portugal, Lisboa), Manuel
Natureza, Ministerio
Benito Crespo (Dto. Ciencias Ambientales y Recursos Naturales, Universidad de
Alicante), Juan Antonio Devesa Alcaraz (Fac. Ciencias, Universidad de Extrema Obon
de Castro (Dto. de Biologa
Vegetal [Botanica],

dura en Badajoz), Concepcion


Vexetal. Fac. Farmacia,
Universidad de Murcia), Santiago Ortiz (Dto. de Bioloxa
Universidad de Santiago de Compostela) and Josep Antoni Rossello` (Dto. de
Vegetal, Universidad de Valencia). Susana Requena Moreno and Manuel
Biologa

1877
Universidad Autonoma

Ruiz (Dto. de Ecologa,


de Madrid) have been of great help
in the statistical process. The advices from Pieter M. Kroonenberg (Dt. of Education, Leiden University) have been particularly useful in the interpretation of the
response of the multiple variable in the sample. Special thanks to Mark W. Schwartz
(Dt. of Enviromental Science and Policy, University of California, Davis) for his
comments of earlier drafts of the manuscript. We also very much appreciate the
reviews from: Christie Brigan, Kelly Lyon, Diane Thompson and Jason Hoeksema
(Dt. of Enviromental Science and Policy, University of California, Davis). This

paper was prepared during a postdoctoral fellowship (Ministerio de Educacion,


Cultura y Deporte / Fulbright, FU2001-50818486) of one of the authors (F.D.L.).
Appendix 1
Taxa included in the statistical analysis.
Allium commutatum Guss.
Allium subhirsutum L.
Anthemis chrysantha J. Gay

Aquilegia pyrenaica subsp. cazorlensis (Heywood) Galiano and Rivas Martnez


Aristolochia bianorii Sennen and Pau
Asphodelus roseus Humb. and Maire
Avena eriantha Durieu
Biscutella dufourii G. Mateo and M.B. Crespo
Mus and Mayol
Brimeura duvigneaudii (L. Llorens) J.A. Rossello,
Carduus paui Devesa and Talavera
Centaurea carratracensis Lange
Centaurea mariana Nyman

Centaurium somedanum Lanz


Cochlearia pyrenaica DC.
Cytisus decumbens (Durande) Spach
Delphinium montanum DC.
Dracocephalum austriacum L.
Epilobium nutans F.W. Schmidt

Erodium paularense Fernandez


Gonzalez
and Izco
Euphorbia gaditana Cosson
Festuca frigida (Hackel) K. Richter
Fumaria flabellata Gasparr.
Genista dorycnifolia Font Quer
Helianthemum alypoides Losa and Rivas Goday
Hieracium glaucophyllum Scheele
Hieracium vegaradanum De Retz
Hyacinthoides vicentina (Hoffmanns. and Link) Rothm. subsp. vicentina
Iris juncea Poiret
Jurinea fontqueri Cuatrec.
and Ortiz

Leucanthemum gallaecicum Rodrguez


Oubina
Limonium carthaginense (Rouy) C.E. Hubb. and Sandwith

Limonium estevei Fernandez


Casas
Limonium migjornense Llorens
Limonium stenophyllum Erben
Linaria huteri Lange
Lysimachia minoricensis J.J. Rodr.
Melilotus italica (L.) Lamk.

Narcissus calcicarpetanus Fernandez


Casas

1878
Appendix 1. (continued)

Narcissus perez-chiscanoi Fernandez


Casas
Nepeta apuleii Ucria ex Guss.
Ononis azcaratei Devesa
Petrocoptis grandiflora Rothm.
Pimpinella procumbens (Boiss.) Pau
Potentilla inclinata Vill.
Puccinellia fasciculata subsp. pungens (Pau) W.E. Hughes
Rosmarinus eriocalix Jordan and Fourr.
Santolina ageratifolia Barnades ex Asso.
Scabiosa andryaefolia (Pau) Devesa
Senecio coincyi Rouy
Sideritis bubanii Font Quer
and Rivera
Sideritis pungens subsp. javalambrensis (Pau) Obon
Sideritis reverchonii Willk.
Silene viridiflora L.
*Tanacetum funkii Schultz Bip. ex Willk.
Teucrium subspinosum Pourr. ex Willd.
Thymelaea granatensis (Pau) Lacaita
Thymus willkommii Ronniger
Vaccinium vitis-idaea L.
Viola crassiuscula Bory
* No autoecological information is known for this taxon. Data for habitat and substrate variables have
been inferred from recorded old locations of the plant.

Appendix 2
Original and recodified variables (level of measure selected appears in brackets) with categories and
frequencies.
Recodified variable a (measure level)

Variable (measure level)


Category

EFFECT OF ANTHROPOGENIC
INTERACTION (Ordinal)
pos
15.25
ndet
50.85
1effec
27.12
2effec
5.08
3effec
1.69
LEVEL OF PROTECTION (Ordinal)
nint
27.12
prot
64.41
maxprot
8.47
TIME OF DESCRIPTION (Ordinal)
clas
59.32
mod
25.42
act
15.25
RATE OF TAXONOMIC SYNONYMY (Ordinal)
nul
33.90
low
47.46
med
8.48
hig
10.17

Category

Effect of anthropogenic
interaction (Ordinal)
pos
ndet
1effect
12effect

15.25
50.85
27.12
6.78

1879

appendix2. (continued)
Recodified variable a (measure level)

Variable (measure level)


Category

LIFE CYCLE (Ordinal)


ann
11.86
bien
3.39
,5
40.68
.5
44.07
TYPE OF DISPERSION UNIT
(Single nominal)b
wind
13.56
villo
15.25
sema
42.37
auto
10.17
endo
8.47
exo
1.69
myrme
6.78
unkn
1.70
TYPE OF POLLINATION (Single nominal)
ento
89.83
ane
5.09
abse
5.09
CLONAL GROWTH (Single nominal)
y
32.20
n
52.54
ndat
15.25
TYPE OF BREEDING SYSTEM
(Single nominal)
facalo
30.51
stralo
18.64
facauto
3.39
aga
8.47
ninfo
38.98
SUBSTRATE b (Single nominal)
calc
45.76
acid
18.64
gyps
3.39
serp
3.39
indi
20.34
sal
8.47
b
HABITAT (Multiple nominal)
wood
6.78
sand
25.42
rock
3.39
weed
11.86
shru
22.03
lago
5.08
clea
8.47
bog
10.17
scree
6.78
BIOGEOGRAPHY (Binomial)
end
71.19
limi
28.81
a

Category

Life cycle (Ordinal)


112

15.25

,5
.5
Type of dispersion unit
(Single nominal)b
wind
villo
sema
auto
zoo

13.56
15.25
42.37
10.17
10.17

myrme

6.78

Type of breeding system


(Single nominal)
facalo
stralo
auto1aga

30.51
18.64
11.86

40.68
44.07

ninfo
Substrate b (Single nominal)
calc
acid
gyps1serp

38.98

indi
sal
Habitat b (Multiple nominal)
wood
ston

20.34
8.47

weed
shru
lago
clea
bog
scree

11.86
22.03
5.08
8.47
10.17
6.78

45.76
18.64
6.78

6.78
28.81

Recodified variables maintain the former names but lose upper case letters; this notation is followed in
tables and plots. bVariables with some lost categories due to recodification; frequencies do not add up to
100%.

1880
Appendix 3
New recodification of Habitat variable and change in measure level of Type of pollination.
Recodified variable a (measure level)

Variable (measure level)


Category

Habitat (Multiple nominal)


wood
5.36
ston
30.36
weed
11.86
scree
7.14
shru
23.21
clea
7.14
lago
3.57
bog
10.71
TYPE OF POLLINATION (Binomial)
ento
94.64
ane
5.36

Category

Habitat (Multiple nominal)


wood
ston
weed
scree
shru
clea
aqua

5.36
30.36
11.86
7.14
23.21
7.14
14.29

Recodified variables maintain the former names but lose upper case letters; this notation is followed in
tables and plots.

References
AAVV 2000. Red list of the Spanish vascular flora (valuation according to IUCN categories). Conser Vegetal 6 (extra): 1138.
vacion
J., Vega J.C., de Hoyos C., Negro A. et al. 1996. Las ultimas

Aldasoro J.J., Aedo C., Munoz


turberas

cantabricas.
Quercus 129: 1619.
Ayensu E.S. and DeFilipps R.A. 1981. Smithsonian Institution endangered flora computerized information. In: Morse L.E. and Henifin M.S. (eds), Rare Plant Conservation: Geographical Data
Organization. New York Botanical Garden, New York, pp. 111122.

de la flora espanola.
Banares
Baudet A. and de Benito J.M. 1990. Estrategia de conservacion
Technical
report. ICONA, Madrid, Spain.

Barreno E., Bramwell D., Cabezudo B., Cardona M.A., Costa M., Fernandez
Casas J. et al. 1984. Listado
Informacion

Ambiental 3: 4871.
de plantas endemicas,
raras o amenazadas de Espana.
Bevill R.L. and Louda S.M. 1999. Comparisons of related rare and common species in the study of plant
rarity. Conservation Biology 13: 493498.
Birks H.J.B. 1987. Recent methological developments in quantitative descriptive biogeography. Annales
Zoologici Fennici 24: 165178.
Briggs J.D. and Leigh J.H. 1996. Rare or Threatened Australian Plants. CSIRO, ANCA, Collingwood,
Australia.
Carroll R., Augspurger C., Dobson A., Franklin J., Orians G., Reid W. et al. 1996. Strengthening the use
of science in achieving the goals of the endangered species act: an assessment by the Ecological
Society of America. Ecological Applications 6: 111.
Carreras J., Carrillo E., Ninot J.M., Soriano I. and Vigo J. 1996. Plantas vasculares del piso alpino de los
Botanico

Pirineos catalanes raras o amenazadas. Anales del Jardn


de Madrid 54: 521527.
Castro Parga I., Moreno Saiz J.C., Humphries C.J. and Williams P.H. 1996. Strengthening the Natural
and National Park system of Iberia to conserve vascular plants. Botanical Journal of the Linnean
Society 121: 189206.

de las lagunas
Cirujano S.,Velayos M., Castilla F. and Gil M. 1992. Criterios botanicos
para la valoracion

y humedales espanoles.
ICONA-CSIC, Madrid, Spain.
Colas B., Olivieri I. and Riba M. 1997. Centaurea corymbosa, a cliff-dwelling species tottering on the
brink of extinction: a demographic and genetic study. Proceedings of the National Academy of
Sciences USA 94: 34713476.
de Lange P.J. and Norton D.A. 1998. Revisiting rarity: a botanical perspective on the meanings of rarity
an the classification of New Zealands uncommon plants. Royal Society of New Zealand, Miscellaneous Series 48: 145160.

1881
Dixon K.W. 1994. Towards integrated conservation of Australian plants the Western Australian model.
Biodiversity and Conservation 3: 148159.

de la flora iberica.

Domnguez
Lozano F. 2000. Herramientas para la conservacion
Aproximacion

historica,
el caso de Vella pseudocytisus ssp. pauiy estudio de los patrones de rareza y amenaza, Ph.D.

Thesis, Universidad Autonoma


de Madrid, Madrid, Spain.
42: 127141.
Fiedler P.L. 1995. Rarity in the California flora: new thoughts on old ideas. Madrono
Anton
M., Maldonado Ruiz J., Morla Juaristi C. and Sainz Ollero H. 2002. Fitogeografa
historica

Garca

de la pennsula
Iberica.
In: Pineda F.D., de Miguel J.M., Casado M.A. and Montalvo J. (eds), La
Pearson Educacion,

Madrid, Spain, pp. 4564.


Diversidad Biologica
de Espana.
Gifi A. 1991. Nonlinear Multivariate Analysis. John Wiley and Sons, Chicester, UK.
Given D.R. and Norton D.A. 1993. A multivariate approach to assessing threat and for priority setting in
threatened species conservation. Biological Conservation 64: 5766.

Gomez
Campo C. 1985. The conservation of Mediterranean plants: principles and problems. In: Gomez
Campo C. (ed.), Plant Conservation in the Mediterranean Area. W. Junk Publishers, Dordrecht, The
Netherlands, pp. 38.

de especies vegetales amenazadas: la experiencia iberica.

Gomez
Campo C. 1990. Metodos
de proteccion

In: Hernandez-Bermejo
J.E., Clemente M. and Heywood V. (eds), Proceedings of the International

Conference on Conservation Techniques in Botanic Gardens, May 1014, 1987, Cordoba.


Koeltz

Scientific Books, Konigstein,


Germany, pp. 3337.

J.M. 1993. Conservation of Iberian endemic plants: the botanical


Gomez
Campo C. and Herranz Sanz
reserve of La Encantada (Villarrobledo, Albacete, Spain). Biological Conservation 64: 155160.
Vegetal. Limusa, Mexico City, Mexico.
Grant V. 1989. Especiacion
Grizzle R.E. 1994. Environmentalism should include human ecological needs. Bioscience 44: 263268.
Hall A.V. 1993. Setting conservation priorities for threatened species: a joint grouping and sequencing
method. South African Journal of Botany 59: 581591.
Hedge S.G. and Ellstran N.C. 1999. Life history differences between rare and common flowering plant
species of California and British Isles. International Journal of Plant Sciences 160: 10831091.

de
Hernandez
Bermejo J.E. and Sainz Ollero H. 1990. Un ndice de estenocoria aplicable a la evaluacion
en diferentes territorios. In: Hernandez-Bermejo

riesgos de extincion
J.E., Clemente M. and Heywood
V. (eds), Proceedings of the International Conference on Conservation Techniques in Botanic

Gardens, May 1014, 1987, Cordoba.


Koeltz Scientific Books, Konigstein,
Germany, pp. 141147.
Herrera J. 1988. Pollination relationships in southern Spanish mediterranean shrublands. Journal of
Ecology 76: 274287.
Hooper S.D. 1998. An Australian perspective on plant conservation biology in practice. In: Fiedler P.L.
and Kareiva P.M. (eds), Conservation Biology for the Coming Decade. Chapman & Hall, New York,
pp. 255278.
Holsinger K.E. and Gottlieb L.D. 1991. Conservation of rare and endangered plants: principles and
prospects. In: Falk D.A. and Holsinger K.E. (eds), Genetics and Conservation of Rare Plants. Oxford
University Press, New York, pp. 195208.
Keith D.A. 1998. An evaluation and modification of world conservation union red list criteria for
classification of extinction risk in vascular plants. Conservation Biology 12: 10761090.
Kroonenberg P.M., Harch B.D., Basford K.E. and Cruickshank A. 1997. Combined analysis of categorical and numerical descriptors of Australian groundnut accessions using nonlinear principal component analysis. Journal of Agricultural, Biological, and Environmental Statistics 2: 294312.
Kruckeberg A.R. and Rabinowitz D. 1985. Biological aspects of endemism in higher plants. Annual
Review of Ecology and Systematics 16: 447479.
Kunin W.E. and Gaston K.J. 1993. The biology of rarity: patterns, causes and consequences. Trends in
Ecology and Evolution 8: 298301.
Lewis D.Q. 1998. A literature review and survey of the status of Iowas terrestrial flora. Journal of the
Iowa Academy of Science 105: 4554.
Lobo J.M., Castro I. and Moreno J.C. 2001. Spatial and environmental determinants of vascular plant
species richness distribution in the Iberian Peninsula and Balearic Islands. Biological Journal of the
Linnean Society 73: 233253.
M. 1994. Monografa
del genero

Luceno
Carex en la pennsula
Iberica
e islas Baleares. Ruizia 14:
5139.

1882

Montserrat P. and Villar L. 1972. El endemismo iberico.


Aspectos ecologicos
y fitotopograficos.
Boletim
da Sociedade Broteriana 46: 503527.
Morse L.E. 1996. Plant rarity and endangerment in North America. In: Falk D.A., Millar C.I. and Orwell
M. (eds), Restoring Diversity. Strategies for Reintroduction of Endangered Plants. Island Press,
Washington, DC, pp. 722.
J., Castro H. and Cabello J. 1996. Agricultural development vs biodiversity conservaMota J.F., Penas
southeastern Spain). Biodiversity
tion: the Mediterranean semiarid vegetation in El Ejido (Almera,
and Conservation 5: 15971617.
Norton B.G. 1986. On the inherent danger of undervaluing species. In: Norton B.G. (ed.), The
Preservation of Species. Princeton University Press, Princeton, New Jersey, pp. 110137.

Palacios C. and Gonzalez-Candelas


F. 1997. Lack of genetic variability in the rare and endangered
Limonium cavanillesii (Plumbaginaceae) using RAPD markers. Molecular Ecology 6: 671675.
Palmer M.A. 1996. A strategic approach to the conservation of plants in the United Kingdom. Journal of
Applied Ecology 33: 12311240.
Palmer M.A., Hodgetts N.G., Wigginton M.J., Ing B. and Stewart N.F. 1997. The application to the
British flora of the world conservation unions revised red list criteria and the significance of red lists
for species conservation. Biological Conservation 82: 219226.
de los Monegros. La paciencia como estrategia de supervivencia. Instituto
Pedrocchi C. 1998. Ecologa
de Estudios Altoaragoneses, Centro de desarrollo de Monegros, Huesca, Spain.

Peinado M., Alcaraz F. and Martnez-Parras


J.M. 1992. Vegetation of southeastern Spain. Flora et
Vegetatio Mundi 10: 1487.
Perring F.H. and Farrell L. 1983. British Red Data Books: 1. Vascular plants. 2nd edn. RSNC, Lincoln,
UK.

Pons A. and Quezel


P. 1985. The history of the flora and vegetation and past and present human

disturbance in the Mediterranean region. In: Gomez


Campo C. (ed.), Plant Conservation in the
Mediterranean Area. W. Junk Publishers, Dordrecht, The Netherlands, pp. 2443.
Rabinowitz D. 1981. Seven forms of rarity. In: Synge H. (ed.), The Biological Aspects of Rare Plant
Conservation. Wiley and Sons, London, pp. 207217.
Ranker T.A. and Arft A.M. 1994. Allopolyploid species and the US Endangered Species Act. Conservation Biology 8: 895897.
Richards A.J. 1997. Plant Breeding Systems. Chapman & Hall, London.
Rojas M. 1992. The species problem and conservation: what are we protecting? Conservation Biology 6:
170178.
Schemske D.W., Husband B.C., Ruckelshaus M.H., Goodwillie C., Parker I.M. and Bishop J.G. 1994.
Evaluating approaches to the conservation of rare and endangered plants. Ecology 75: 584606.

Zarzoso J.C. 1994. La flora vascular espanola:


Ecologa
8: 203225.
Simon
diversidad y conservacion.
Skinner M.W., Tibor D.P., Bittman R.L., Ertter B., Ross T.S., Boyd S. et al. 1995. Research needs for
42: 211241.
conserving Californias rare plants. Madrono
Soltis P.S. and Gitzendanner M.A. 1999. Molecular systematics and the conservation of rare species.
Conservation Biology 13: 471483.
SPSS Inc. 1994. SPSS Categories 6.1. SPSS, Chicago, Illinois.
Thompson J.D. and Lumaret R. 1992. The evolutionary dynamics of polyploid plants: origins, establishment and persistence. Trends in Ecology and Evolution 7: 302307.

Toledo M.V. 1993. La riqueza florstica


de Mexico:
un analisis
para conservacionistas. In: Guevara S.,
Moreno-Casasola P. and Rzedowski J. (eds), Logros y perspectivas del conocimiento de los recursos

Sociedad Botanica

vegetales de Mexico
en vsperas
del siglo XXI. Instituto de Ecologa,
de Mexico,
Mexico, pp. 109123.

S. and Jorgensen P.M. (eds) 2000. Libro rojo de las plantas


Valencia R., Pitman N., Leon-Yanez

endemicas
del Ecuador 2000. Herbario QCA, Pontificia Universidad Catolica
del Ecuador, Quito,
Equador.
Weller S.G. 1994. The relationship of rarity to plant reproductive biology. In: Bowles M.L. and Whelan
C.J. (eds), Restoration of Endangered Species. Conceptual Issues, Planning and Implementation.
Cambridge University Press, Cambridge, UK, pp. 90117.
Whitman T.G., Morrow P.A. and Bradley M.P. 1991. Conservation of hybrid plants. Science 254:
779780.