Beruflich Dokumente
Kultur Dokumente
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
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;
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.
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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
Results
The PRINCALS algorithm calculates parameters in a similar manner to other
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Table 1. Variables and categories (identification labels used in later statistics appear in brackets).
Variable
Categories
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 POLLINATION
CLONAL GROWTH
SUBSTRATE
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Table 1. (continued)
Variable
Categories
HABITAT
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
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.
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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-
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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
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.
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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.
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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
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Table 4. Protection measures for the second group of taxa.
Taxa
Protected area
Legal cover
Habitat Directive. Asturian Catalogue of
Threatened Species
Catalonian plan of natural interest sites
Andalucian Catalogue of protected 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
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
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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
1877
Universidad Autonoma
1878
Appendix 1. (continued)
Appendix 2
Original and recodified variables (level of measure selected appears in brackets) with categories and
frequencies.
Recodified variable a (measure level)
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)
Category
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
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)
Category
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.
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